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Published by Field Museum of Natural History V^ / &*-'

VOLUME 37

A TAXONOMIC REVISION OF THE

GENUS CNEMIDARIA (CYATHEACEAE)

ROBERT G. STOLZE

October 28, 1974

FIELDIANA: BOTANY

A Continuation of the

BOTANICAL SERIES

of

FIELD MUSEUM OF NATURAL HISTORY

VOLUME 37

FIELD MUSEUM OF NATURAL HISTORY

CHICAGO, U. S. A.

A TAXONOMIC REVISION OF THE

GENUS CNEMIDARIA (CYATHEACEAE)

FIELDIANA

Botany

Published by Field Museum of Natural History

VOLUME 37

A TAXONOMIC REVISION OF THE

GENUS CNEMIDARIA (CYATHEACEAE)

ROBERT G. STOLZE

Custodian, Pteridophyte Herbarium

Field Museum of Natural History

October 28, 1974

Publication 1194

PATRICIA M. WILLIAMS

Managing Editor, Scientific Publications

Library of Congress Catalog Card Number: 74-791 75

US ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

INTRODUCTION

Cnemidaria, a genus of tropical American ferns belonging to the tree-

fern family, Cyatheaceae, is confined largely to wet habitats, hi or at

the edges of forests, at elevations ranging from sea level to 2,300 m.

(typically 500-2,000 m.). These ferns are mostly subarborescent, with

leaves caespitose, their caudices rarely attaining a thickness of more than

7 cm. or a length of more than a meter. Individual leaves, however, may

reach great proportions, occasionally up to a length of 3.5 m. in some

Antillean species. Within the family the species of Cnemidaria are

characterized by the generally acaulescent habit, a comparatively simple

leaf architecture (never more than pinnate-pinnatisect), the basal vein-

lets conniving strongly or forming costal areoles, a general lack of tri-

chomes on the axes, and the spores possessing large, regularly disposed

pores. A total of 23 species is recognized in this treatment.

In his Tentamen Pteridographiae (1836), C. B. Presl proposed the

genus Cnemidaria, based on five species formerly included within

Cyathea and Hemitelia. These species he considered distinct, basing

his decision chiefly on the character of the costal areoles formed by

anastomosing basal veins. Later (1847) he created three more genera,

Actinophlebia, Hemistegia, and Microstegnus, into which he placed

most of his original species of Cnemidaria along with several other taxa.

In Cnemidaria itself he maintained only one species: C. speciosa. Sub-

sequent authors, (Kunze, Fee, William Hooker, et al.} chose to treat

Presl's original species of Cnemidaria under Hemistegia and (or)

Hemitelia, and for over 50 years the genera of Cyatheaceae were sep-

arated primarily by the presence, absence or shape of the indusium.

Maxon (1912) considered that a number of species of Hemitelia

formed a natural group and brought them together under the subgenus

Cnemidaria. In this subgenus were included all the earlier "genera" of

Presl, as well as a number of new species, many of which were based on.

single collections. They were distinguished from the subgenus "Euhemi-

telia" either by the presence of costal areoles, or by the "scarcely

arborescent" habit and less-dissected leafy parts. Maxon's work cleared

up much of the earlier confusion in the nomenclature and in the typifica-

tion of species.

1

2 FIELDIANA: BOTANY, VOLUME 37

Copeland (1947) concurred with Maxon on the naturalness of the

group, and restored Cnemidaria to generic rank. This concept was sup-

ported by Holttum & Sen (1961), who reinforced the distinctiveness of

the genus by pointing out the peculiar character of the spores. They

further suggested that some of the free-veined species earlier placed in

the genus lacked this type of spore and thus should be excluded. Tryon

(1970) included Cnemidaria in his classification of the family, refined

the generic concept, and pointed out still another pertinent character to

further delimit the genus within the Cyatheaceae: that trichomes are rare

or lacking on the adaxial side of the costa. Other genera are characterized

by the common occurrence of trichomes along most of the axes, par-

ticularly on the adaxial side.

Throughout the years, poor or incomplete collecting has contributed

to lack of understanding of the generic features of tree ferns. Due to the

size of individual plants, collectors either ignore them in the field or are

satisfied with a pinna or two. The problem remains a current one. How-

ever, enough material (nearly 2,000 specimens) has been gathered for

this study to indicate that the group of species as treated here forms

perhaps the most natural and distinctive genus in the family. In addition

to the characters already noted in this "Introduction," Cnemidaria also

has whitish or bicolorous petiole scales, rudimentary paraphyses, arach-

noid scurf on the axes, hemitelioid indusia, and pinnae never articulated

at the base. These characters, however, may also occur in species of other

genera.

EVOLUTION AND GEOGRAPHY

The principal characters of Cnemidaria rarely, if ever, occur in other

genera of Cyatheaceae. Within the family, only Cnemidaria has spores

with large, regularly disposed, equatorial pores. Furthermore, the basal

veins of nearly all of the species either merge to form costal areoles or

are at least strongly connivent to the sinus, whereas veins of the species

of the other genera are normally free and non-connivent. Minute,

terete, recurved trichomes are found on relatively few species of Cnemi-

daria, and then generally on the abaxial side of the axes, although this

kind of trichome is characteristically abundant along the axes throughout

the other genera. It is important to note that these trichomes are present

on the adaxial side of the rachis only in the free-veined species of

Cnemidaria, which thus links two primitive characters together in the

same few species. Finally, the leaf architecture in the genus shows less

dissection (never fully bipinnate) than in the remainder of the scaly

Cyatheaceae, whose leaves are typically twice- to thrice-pinnate.

So it appears that Cnemidaria is the most advanced genus of the

family, with more complex venation and simpler lamina architecture

being the end products of its adaptive development. A majority of the

species are highly specialized in these characters. Such evolutionary

developments also occur in at least one species of Trichipteris and in

several species of Cyathea. T. williamsii of Panama is not only simply

pinnate but has costal areoles as well; and in Cyathea there are several

species exhibiting costal areoles, e.g., C. conformis, C. woronovii, and

C. panamensis, as well as a few with subentire pinnae, e.g., C. integrifolia

and C. speciosa Humb. & Bonpl. ex Willd. (not Cnemidaria speciosa

Presl). All of the above species of Cyathea have a partially developed

(hemitelioid) indusium. This character is common to Cnemidaria and,

with the other similarities mentioned, indicates that Cnemidaria has

probably arisen from this particular line of Cyathea. Tryon (1970) has

also indicated the derivation of Cnemidaria from Cyathea.

Within the genus, a phylogeny (fig. 1) can be derived from the oc-

currence of advanced or primitive characters in the species. The most

primitive species is Cnemidaria amabilis, which has the greatest number

of unspecialized features (see discussion under that species) and ex-

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STOLZE: REVISION OF CNEMIDARIA 5

hibits the closest links with Cyathea, especially in venation and indument.

Other primitive species are those, such as Cnemidaria singularis and

C. tryoniana, with free veins and deeply dissected pinnae. The free-

veined C. mutica might represent an ancestral type that gave rise to

species with costal areoles, such as C. horrida, which is probably the

most primitive of the predominately South American areolate group.

The most advanced species are C. karsteniana and C. nervosa, which

are strongly areolate, simply pinnate, and virtually glabrous and acaules-

cent.

The geography of at least one group of species can be seen to corre-

late with its line of evolution, and this suggests the route of migration

along which speciation has occurred. The evolutionary line connects the

very primitive C. amabilis in Venezuela with the simple-veined but less

primitive C. mutica in Panama and Costa Rica, with especially strong ties

to var. chiricana. The line then passes, in Costa Rica, to C. choricarpa

which has costal areoles and less deeply-cut pinnae, but still closely re-

sembles C. mutica var. mutica in many respects. Continuing northward

into Guatemala and Mexico, the evolutionary line reaches an advanced

state in C. decurrens, a relative of C. choricarpa with subentire pinnae.

Other evolutionary lines may have developed in a similar geographic

manner, but their species are now often allopatric. One such line, origin-

ating with C. horrida, passes from C. spectabilis to C. consimilis, and

finally to C. karsteniana, a species endemic in Venezuela. Another line

passes from C. spectabilis through C. speciosa and C. ewanii, and thence

to C. nervosa of Ecuador and Peru. In both lines, the progression is from

species with deeply dissected pinnae and considerable indument on the

axes, to species nearly lacking in indument and with pinnae entire or

subentire.

The greatest number of species, as well as the most primitive ones,

are principally near the center of the range, whereas a majority of the

highly advanced species are near the borders of the range (fig. 1, map 1).

Hence the most likely place of origin of Cnemidaria could be construed

as that area in which the most primitive species are currently found, and

in which there is the greatest present concentration of species. Thus it is

likely that the genus originated in the area of Colombia and northern

Venezuela, for here are found a total of 1 1 species, including the three

most primitive.

A fossil record is reported by Graham and Jarzen (1969) from an

Oligocene formation at San Sebastian, Puerto Rico, and, although

positive identification is difficult from the photomicrograph, it strongly

resembles a Cnemidaria spore not unlike those of C. horrida, a species

FIELDIANA: BOTANY, VOLUME 37

MAP 1. Generalized range of Cnemidaria (outline) and area of highest con-

centration of species (black): 11 species occur in this area, including the three

most primitive ones: C. amahilis, C. sinful fin's, and C. tryoniana.

found currently in Puerto Rico. Graham and Jarzen also mention similar

spores from a Miocene formation in Veracruz, Mexico. There is another

report by Hammen and Gonzalez (1960) of fossil spores from Upper

Pleistocene and Holocene formations in Colombia, and the excellent

photomicrograph shows these to be unquestionably Cnemidaria. How-

ever, these records are too few at present to be useful in determining the

area of origin of the genus.

The distribution of Cnemidaria is generally continuous, with species

overlapping through most of the range. However, there are two notable

regions of disjunction: southeastern Brazil, and an area containing the

provinces of Chiapas and Oaxaca in Mexico and Alta Verapaz in

Guatemala. There is no record of the genus between C. decurrens in

Guatemala and C. choricarpa in Costa Rica, even though that area has

been rather well-collected in recent years, and there is a much larger

gap (2,000 km.) between the collections of C. uleana in Brazil and

C. speciosa in Bolivia. Such disjunctions are apparently due to lack of

proper growing conditions in the intervening regions. Cnemidaria is typi-

STOLZE: REVISION OF CNEMIDARIA 1

cally found on slopes in or at the edges of forests, at middle elevations

in wet areas where there are no prolonged dry periods. A lack of any

one of these factors will apparently result in an unfavorable habitat.

A list of the distribution of species according to country or area

follows :

MEXICO: apiculata, decurrens.

GUATEMALA: decurrens.

COSTA RICA: choricarpa, horrida, mutica vars. chiricana, contigua, .grandis,

and mutica.

PANAMA: cocleana, mutica vars. chiricana, grandis, and mutica.

GREATER ANTILLES: horrida.

LESSER ANTILLES: grandifolia vars. grandifolia and obtusa.

TRINIDAD: consimilis, grandifolia var. obtusa, spectabilis var. spectabilis.

TOBAGO: grandifolia var. obtusa, spectabilis var. spectabilis.

FRENCH GUIANA: cruciata, spectabilis var. spectabilis.

SURINAM: spectabilis var. spectabilis.

BRITISH GUIANA: roraimensis, spectabilis var. spectabilis.

VENEZUELA: amabilis, consimilis, grandifolia var. obtusa, horrida, kar-

steniana, spectabilis var. spectabilis.

COLOMBIA: chocoensis, choricarpa, ewanii, horrida, quitensis, singularis,

spectabilis var. colombiensis, tryoniana, uleana var. abitaguensis.

ECUADOR: ewanii, horrida, nervosa, quitensis, uleana var. abitaguensis.

PERU: alatissima, horrida, nervosa, speciosa, uleana var. uleana.

BOLIVIA: speciosa.

BRAZIL: uleana var. uleana.

MORPHOLOGY

In spite of the large number of specimens examined in this study, some

morphological data are still incomplete, especially pertaining to those

species represented by only a few collections. Collectors often neglect

to gather adequate material for tree fern specimens, either because they

feel it is time-consuming or because they fail to appreciate the taxonomic

problems involved. Like any other fern, tree ferns can be properly classi-

fied only on the basis of characters of the entire plant, and a specimen

without petiole or leaf apex may lack the very portion by which it is

readily distinguished from related species. Thus a specimen of any large

fern, to be of optimum value, should contain leaf apex, a section of rachis

with several central pinnae, a few of the lowermost pinnae, and a basal

section of petiole. A small portion of the caudex (or perhaps a cross-

section of a very thick one) is also desirable. Where this is impractical,

adequate label data should be included, noting length, diameter, type

of scale, spines, etc. With little extra time and effort, a complete sample

of the plant can be adequately collected for mounting on two herbarium

sheets.

A discussion of morphological features is presented here, with special

emphasis on those characters which are particularly diagnostic in

Cnemidaria. Explanation is given for terminology used throughout the

text and in the key.

CAUDEX

Among the species, caudices vary in length up to 1.5 m. and in dia-

meter up to 7 cm. However, relatively few collections include this por-

tion of the plant, or even label data describing it. Where such data are

available, they appear to be of little taxonomic value at the species level.

PETIOLE AND RACHIS

The primary axis of the leaf in species of Cnemidaria is terete on the

abaxial side, shallowly or deeply sulcate on the adaxial side,1 with color

varying from dark brown to light brown to stramineous on dried speci-

1 Throughout the descriptions and key, in the interest of brevity, the "adaxial

side" will be referred to as "above" and "abaxial side" as "beneath."

STOLZE: REVISION OF CNEMIDARIA 9

mens. Toward the apex the rachis is always light colored, but the shade

commonly deepens toward the base of the petiole. Color may vary

among plants of a given species. Frequency and color of scales on the

FIG. 2.

35 (GH).

Portion of leaf showing alate rachis: Cnemidaria choricarpa, Nisman

petiole and rachis can be extremely valuable in identifying species, and

their importance is discussed under "Indument" below.

The rachis is generally smooth in Cnemidaria, but minute spines can

be found, particularly toward the base, in C. consimilis, C. horrida, C.

quitensis, and C. spectabilis. The petiole may be smooth or muricate

or quite spiny, and this may be inconstant even within a variety (e.g.,

C. mutica var. mutica}. But presence or lack of spines can be diagnostic

in several species, especially when used in conjunction with other charac-

ters.

The narrow, green wing of tissue along each side of the rachis (fig. 2)

can be a valuable character. Although lacking in most species and oc-

curring sporadically in others, it is particularly prominent in Cnemidaria

alatissima, C. choricarpa, and C. decurrens. In some species, a scarcely

observable wing may run downward from the decurrent base of the

apical section, and extend as far as the second or third pair of pinnae.

In C. choricarpa the rachis is generally alate for most of its length, and in

C. alatissima, the wing is prominent well down the petiole. In C. decur-

rens the wing is often worn away so that in older plants only the rem-

nants, or merely the lines marking the previous points of attachment may

be seen. On many mounted specimens, especially those having a con-

siderably thickened rachis, the wing may be difficult to detect from

10

FIELDI AN A: BOTANY, VOLUME 37

FIG. 3. Types of leaf apex: a, no distinct apical section, Cnemidaria cruciata,

LePrieur s.n. 1838 (US); b, nonconform apical section, C. spectabilis var. spec-

tabilis, Hombersley 30 (US).

beneath. Therefore in mounted material it is often necessary to view

the rachis at an oblique angle to be certain there is a wing present. On

specimens mounted with adaxial side up, the wing, if present, will be

generally obvious to the naked eye.

One character which merits brief mention is that of the pneumato-

phores (or pneumathodes), which occur along each side of the petiole.

These are areas of soft surface tissue which evidently function in gas

exchange with the inner tissues. In fresh material they may be seen as

a single (sometimes double) line of linear to lanceolate spindle-shaped

areas, pale or light green in color as contrasted with the darker color

of the petiole tissue. In dried material these pneumatophores may be

difficult or impossible to detect, and hence of questionable taxonomic

value at present. However, future study of their size, shape, and spacing,

STOLZE: REVISION OF CNEMIDARIA

11

FIG. 3. c, subconform terminal pinna, with elongated basal lobes, C. speciosa,

Schunke A-143 (US); d, conform terminal pinna, C. ewanii, Cuatrecasas 9022

(US).

especially in living plants, may provide further clues to the relationship

of species.

LAMINA

The lamina in Cnemidaria is broadly lanceolate to ovate-oblong, and

in larger species may attain a length of more than 2 m. In fully grown

specimens the pinnae may number 20 to 30 pairs, rarely less than a

dozen. However, one unusual species, C. singularis, may be distinguished

from all others by its few (three to six pairs) pinnae.

Within the genus the basal pair of pinnae are usually considerably

shorter than the penultimate pair, and also somewhat deflexed (some-

times as much as 45°). This feature can vary substantially from plant

to plant, but at least one, C. tryoniana, can be separated from similar

12 FIELDIANA: BOTANY, VOLUME 37

species partly on the fact that its basal pinnae are scarcely or not at all

reduced.

Within other genera of the Cyatheaceae pinnae are commonly long-

stalked and articulate to the rachis. Pinnae in Cnemidaria are never ar-

ticulate to the rachis, and are commonly sessile or subsessile. An excep-

tion is found in C. bella (fig. 16), where the larger pinnae may have

stalks to 2 cm. long; hence the character is diagnostic. This unusual

character is part of the evidence suggesting that it may be a hybrid (see

also discussion of the species), and hence is atypical of Cnemidaria.

A high degree of variation may be encountered in depth of dissection

of pinnae and in the shape of ultimate segments. Therefore, these charac-

ters may not be particularly useful in separating certain species. Indeed,

they may be confusing. Where dissection of the pinnae is used as a key

character, the basal pair of pinnae and reduced apical ones are excluded.

Apical and basal pinnae, due to their marked reduction in size, generally

have their segments differing from other pinnae in length and shape of

the segments. Thus, in a leaf where central pinnae are cut three-fourths

to seven-eighths to the costa, and the segments are acute to acuminate,

a reduced apical or basal pinna may be cut less than halfway to the

costa and its segments obtuse. However the character, when properly

utilized, can be extremely valuable in delimiting certain groups of

species, or individual species. In the extremely variable C. mutica (see

fig. 13), it is useful at varietal rank, and a detailed discussion is pre-

sented under that species.

A peculiar feature is present in the pinnae of C. cruciata, C. specta-

bilis var. colombiensis , and C. quitensis (fig. 18). The segments are

commonly enlarged toward the tips and curved in such a way as to

cause the margins to crowd or overlap those of adjacent segments. Com-

comitantly in these species there is such a strong convergence of veins

at the sinus that a thickened, cartilaginous mass is formed. In dried

specimens of C. quitensis and C. spectabilis var. colombiensis the sinuses,

particularly as viewed from above, exhibit a light yellowish or brownish

color which contrasts markedly with the adjacent tissue. In other species,

the segments commonly diverge at a broader angle, and only a few

veins converge at the sinus, where the tissue is uniformly colored.

The apex of the lamina can be especially diagnostic in delimiting

groups of species within Cnemidaria (fig. 3). Throughout the key refer-

ence is made to "conform," "subconform," or "nonconform" apical

pinnae. Although it may not be morphologically correct to describe the

terminal segment of the lamina as a pinna, I have used this terminology

as a matter of convenience and conciseness. A conform terminal pinna

STOLZE: REVISION OF CNEMIDARIA

13

5- «W V; >^l&£feg^

M - \'j ^r -^! . f^,ij^s^--

5mm

Cyathaa multiflora

FIG. 4. Trichomes; adaxial side of rachis, costa, and base of costule (insert =

6.5 mm. diam.): Cyathea multiflora, Burger & Malta 4237 (F).

is an apical section which closely corresponds in size and shape to typical

pinnae on the leaf, usually resembling those of the second and third pair.

Subconform is a term given to an apical section which is otherwise con-

form but has a pair of elongated or oversized basal lobes. A nonconform

terminal pinna is a distinct apical section which differs from the typical

pinna in shape and width (and often length). Its general outline is

broadly triangular, widening gradually from apex to base, and having

especially elongated basal lobes.

The most unusual types of leaf apex occur in C. cruciata and C. singu-

laris. Leaves of the former (fig. 3a) completely lack a distinct apical

section, as the pinnae gradually diminish in size all the way to the tip.

In C. singularis (fig. 15) nearly half the lamina appears to be an apical

section, which actually dwarfs each of the main pinnae.

INDUMENT

In Cnemidaria the indument is confined to the petiole, rachis, other

axes, and veins, whereas the laminar tissue is glabrous. Character and

color of the scales may vary on a plant depending upon their location on

the leaf, and provide an important means of identifying species. Tri-

14

FIELDIANA: BOTANY, VOLUME 37

6cm

2cml

FIG. 5. Scales: bicolorous petiole scales, a, Cnemidaria spectabilis var. specta-

bilis and b, C. consimilis; white petiole scales, c, C. amabilis; d, typical amorphous

costal scales; bullate costule scales, e, C. grandifolia.

chomes are of one basic type, and while their value as a diagnostic

character applies only to a few species, their relative position on the

axes is of great significance in delimiting the genus from others in the

Cyatheaceae (see "Evolution and Geography" above). A third type

of indument is a whitish, cobwebby tomentum often thickly matted

on any of the axes, which has been referred to as "arachnoid scurf." It

is of little taxonomic value within the genus, but seems to be confined

to species of Cnemidaria and to a few rather closely related species in

Cyathea.

The trichomes are identical with a type that is characteristically found

in other genera of the Cyatheaceae (fig. 4). They are minute (±0.3

mm.), stiff, terete, recurved, pluricellular and relatively colorless. In

many species of Cnemidaria they are virtually absent. On others they

may be found scattered sparsely along the rachis and costae beneath.

On a few primitive species they are quite abundant on most of the

axes beneath, and even on the rachis above. However, they are always

lacking on costae and costules above — areas where they are most

copious in species of other genera. Species on which trichomes may be

STOLZE: REVISION OF CNEMIDARIA

15

5mm

FIG. 6. Arachnoid scurf: abaxial side of rachis and costa (insert = 1.5 mm.

diam.); Cnemidaria mutica var. mutica, Nisman 22 (GH).

particularly numerous are C. amabilis, C. quitensis, and varieties of C.

mutica except var. contigua.

The color and placement of petiole scales serve to distinguish many

species. Those of C. spectabilis var. spectabilis (fig. 5a), for example,

are located at the very base, and are deep brown with narrow whitish

margins. This particular pattern in bicolorous petiole scales is character-

istic of most species. Bicolorous scales occur along most of the petiole

length in C. consimilis (fig. 5b), but the brown median stripe is com-

monly narrower than either of the whitish margins. Petiole scales of this

pattern also occur on C. ewanii, C. quitensis, C. speciosa, and C. uleana.

During the course of this study an attempt was made to classify a num-

ber of species on the basis of this distinction, i.e., petiole scales; a) con-

colorous; b) bicolorous, with median stripe much broader than either

of the white margins; c) bicolorous, with median stripe much narrower

than either margin. On material examined, scale color and pattern

seemed constant within each species, but a sufficient number of petioles

was not available for the results to be conclusive. The character, how-

16

FIELDIANA: BOTANY, VOLUME 37

FIG. 7. a, Basal veins anastomosing, Cnemidaria grandifolia var. .grandifolia,

Bailey 274 (GH); b, basal veins free, but connivent, C. mutica var. mutica, Nis-

man62 (GH).

ever, merits future consideration. Scales on the petioles of C. amabilis

(fig. 5c) and C. alatissima are white (rarely with minute splotches of

brown), lacking a brown median stripe. On both species, the color is

constant well up onto the rachis, so that even specimens lacking

petioles may be distinguished on this basis, and it is a most useful key

character.

Shape of scales may be relatively constant on various parts of the

plant, e.g., linear, linear-lanceolate, or lance-ovate on the petiole, bullate

(in one species) with hair tips on the costule, but the most predominant

scale along all the upper axes in Cnemidaria is characterized by its lack

of definite shape (fig. 5d). Scales scattered along the rachis, costae, and

costules of most species are generally broad, with irregular margins, and

are appressed to the axis. These may be white, brown, or bicolorous.

The most distinctive of all Cnemidaria scales occur only in C. grandi-

folia, along the costules beneath, and are described as "bullate" (fig. 5e).

These are blunt or, more frequently, with long-attenuate hairlike tips,

and have each margin folded around to touch or overlap the other, which

gives them an inflated appearance. This is one of the characters which

effectively separate this species from C. horrida, whose costular scales,

when present, are of the typical flattened, amorphous type. Bullate scales

are quite common throughout other genera of Cyatheaceae.

Arachnoid scurf (fig. 6), usually appearing macroscopically as faint

patches of whitish powder, can be a variable character in Cnemidaria as

to position, frequency, and abundance on the axes. It can also be in-

constant in its nature: always whitish and only one cell in thickness, but

sometimes appearing in thick, scalelike masses, or at other times like

filamentous trichomes. Occasionally it may be scalelike at the point of

FIG. 8. Branching of basal acroscopic veins: a, veins once-forked, Cnemidaria

mutica var. chiricana, Killip 5350, (US); b, veins simple, C. amabilis, Steyer-

mark 94886 (NY).

FIG. 9. Position of basal basiscopic veins: a, arising from costule, Cnemidaria

mutica var. mutica, Scamman & Holdridge 7881 (GH); b, arising from the

costa, C. uleana var. uleana, Tryon & Tryon 687 1 (GH).

17

18 FIELDIANA: BOTANY, VOLUME 37

attachment, but tapering to an extremely long hairlike tip, one cell broad,

but many cells long. In these instances, it appears to be a modified scale

in its origin. When hairlike, they have no similarity to the relatively

thickened, terete trichomes discussed above. Instead, they can be

long and tortuous, with their cells flattened and extremely elongated.

Arachnoid scurf can be quite abundant on the axes of C. horrida or C.

mutica, at times obscuring portions of the rachis or costa, or it may be

completely lacking on species such as C. amabilis and C. uleana. On

some specimens it may be confined almost entirely to the point of junc-

ture of two axes. Of little or no apparent taxonomic significance within

the genus, it may still be considered as a generic characteristic of

Cnemidaria, and as atypical in other genera of Cyatheaceae.

VEINS

A characteristic feature in Cnemidaria is the anastomosing of basal

veins to form costal areoles. It has been discussed above (see "Evolution

and Geography") as one of the important clues to phylogeny within the

genus as well as to evolutionary development of Cnemidaria within the

family. In a majority of the 23 species, the basal vein in each segment

forks once or several times, and the lowermost branches of adjacent

veins are joined by a transverse veinlet, thus forming an areole along

the costa (fig. 7a). Veins of several species lack such a connecting

veinlet, and whereas the basal veins are directed toward those of the

adjacent segment, they bend rather sharply and are merely connivent to

the sinus (fig. 7b), therefore no areoles are formed. In only one species,

C. amabilis, the veins are scarcely or not at all connivent, and they

reach the segment margin at a point just above the sinus (fig. 8b). This

condition is typical of the venation of the other genera of Cyatheaceae.

The degree of branching of veins is rather variable within the genus

and is usually unreliable as a diagnostic character. Veins in some species

may be 0- to 3-forked, or in others 2- to 4-forked. Often this appears

correlated with the size of the particular leaf. However, in all but a few

species the basal veins are always at least once-forked (fig. 8a). One

exception is C. amabilis, in which the basal acroscopic veins are always

simple, which thus provides a useful means of separating this species

from the closely-related C. mutica var. chiricana. Two other species

which might be confused may also be distinguished partly on the basis

of venation: C. karsteniana has veins mostly once-forked, whereas the

veins of C. nervosa are always simple.

The position of basal veins on the axes is another character which

may be somewhat variable on some species, yet distinctive in others. In

STOLZE: REVISION OF CNEMIDARIA

19

FIG. 10. Position of soral lines: a, submarginal, Cnemidaria apiculata, Hall-

berg 1557 (NY); b, supramedial, C. spectabilis var. spectabilis, Miller & John-

son 164 (F); c, inframedial, C. tryoniana, Metcalf & Cuatrecasas 30122 (GH).

the typical condition the lowermost veins on either side of the segment

arise from the costule at or near its base (fig. 9a). However, in several

species the basal basiscopic veins commonly arise from the costa (fig.

9b), e.g., Cnemidaria. chocoensis, C. singularis, C. tryoniana, and C.

uleana. Where this character is used in the key to separate the latter

two from C. mutica, the condition is best observed in the basal third or

apical third of the pinna. At these points the basal basiscopic veins are

constant in their position on the costa, whereas toward the center of the

pinna they may assume the more typical position at the base of the

costule.

FIG. 11. Shape of indusium: a, spathulate, Cnemidaria apiculata, Jurgensen

873 (NY); b, semicircular, C. horrida, Lopez 143 (US); c, circular, C. cocleana,

Tyson et al. 2452, (FSU).

20 FIELDIANA: BOTANY, VOLUME 37

SORI AND INDUSIA

In Cnemidaria, sori are more or less globose and placed on the veins

(when branched) generally above the fork. On some larger species the

sori may be arranged in several series, but typically they are disposed

in a single line between the costule and segment margin (fig. 10). On

fully mature plants the soral pattern may be so regular and unbroken

as to form a continuous zigzag line along each side of the costa, from

base to apex of pinna. On certain species, such as C. mutica, the sori

may vary in their relative position on the segment — placed nearer the

costule on some plants, or nearer the margin on others — so that within

the species the soral line may be inframedial, medial, or supramedial.

In other species the relative position of the sorus is constant and can

be used effectively as a diagnostic character. In C. bella and C. apicu-

lata (fig. lOa) the soral line is placed so close to the edge of the segment

as to appear almost marginal. In C. spectabilis var. spectabilis the sori

are located away from the margin, and yet are nearer the margin than

the costule (fig. lOb). With other species, e.g., C. quitensis and C. tryon-

iana (fig. lOc), the soral line is inframedial, at times nearly touching

the costule; in this case the veins are usually branched at or near the

costule, and the sori placed at or just above the forks.

The indusium in Cnemidaria is characteristically hemitelioid, i.e., it

subtends the sorus, partially surrounding the receptacle (fig. lib). It is

roughly semicircular (commonly 120-180°), saucer-shaped, with mar-

gins subentire, erose or 2- to 3-lobed. In young plants of C. horrida and

a few other large species some of the indusia may be subcyatheoid, i.e.,

irregularly cup-shaped to subglobose, thus almost covering the sorus.

In respect to the degree the indusium encircles the receptacle, there are

a few notable exceptions within the genus. In C. apiculata the indusia are

very narrow and spathulate (fig. lla), or in young plants sometimes so

strongly cupped as to appear narrowly cucullate. In C. nervosa and

C. cocleana, the indusium completely encircles the sorus (fig. lie). This

character is so unique in these three species that it may be used exclu-

sively in the key to distinguish them from others.

SPORES

Cnemidaria spores are unique in the family. Features of size, shape,

and surface ornamentation are not particularly significant, but the large

pores located on each side constitute an excellent generic character.

Mature spores are trilete and have an approximate lateral diameter of

37-52^. Ferine is often lacking or very thin and closely surrounding the

exine, and there are no definable surface patterns. However, in all

STOLZE: REVISION OF CNEMIDARIA

21

S E M MICROGRAPHS

(X 2 0 0 0)

PROXIMAL

DISTAL

CYATHEA MULTIFLORA

CYATHEA SPECIOSA

CNEMIDARIA SPECTABILIS

FIG. 12. SEM micrographs (c. 2000 X ) of Cyatheaceae spores: without large

equatorial pores, a, Cyathea multiflora, b, C. speciosa; with typical large equa-

torial pores, c, Cnemidaria spectabilis.

Cnemidaria spores there is one large pore (up to 15 /* in diameter) lo-

cated near the center of each side, on or near the equator (fig. 12c).

In Erdtman and Sorsa (1971) this aperture is referred to as "a

conspicuous lenticular hollow or depression in the exine." These are

easily visible under a light microscope, but the results are much more

dramatic when examined by means of a scanning electron microscope.

Such pores are lacking in spores of all other genera of the Cyatheaceae.

Cnemidaria spores may also have small apertures or depressions,

quite irregular in size and spacing, scattered along the surface, and this

feature is sometimes also seen in the spores of other genera. For example,

Cyathea spores (figs. 12a, b) may have hundreds of minute apertures

covering the surface, arranged at random or in vague patterns, although

in this instance they are much more constant in size and shape than those

of Cnemidaria. A species considered by Maxon (1912) as part of the

subgenus Cnemidaria was Hemitelia speciosa (Willd.) Kaulf. (= Cya-

thea speciosa Willd.). These plants resemble Cnemidaria in that their

pinnae are subentire, but they do not conform in characteristics of

venation and indument. Examination of the spores (fig. 12b) confirms

the evidence that this species does not belong in Cnemidaria.

22 FIELDIANA: BOTANY, VOLUME 37

Scanning electron micrographs are presented on pages 78-79 to illus-

trate spores of a number of species of Cnemidaria. To prepare spores

for SEM photography, aluminum stubs were first coated with silver con-

ductive paint and allowed to dry. Sporangia were then crushed in a

drop of distilled water on the stubs and bits of sporangia! walls removed

from the surface. After drying, stubs were vapor coated with 200 A

gold and examined under a Cambridge Stereoscan Mark II-A SEM.

MATERIALS

This revision is based on the study of nearly 2,000 specimens which

were either loaned to me by various institutions or made available to

me on visits to their herbaria. I also made several visits to Gray Herbar-

ium to examine specimens on loan to Dr. Rolla Tryon from Berlin,

Paris, Stockholm, Caracas, and the Missouri Botanical Garden. These

herbaria, with the abbreviations of Lanjouw and Stafleu, Index Her-

bariorum, 1964, are listed here:

B — Botanisches Museum, Berlin, Germany

C — Botanical Museum and Herbarium, Copenhagen, Denmark

CR — Museo Nacional de Costa Rica, San Jose, Costa Rica

F — Field Museum of Natural History, Chicago, Illinois

FSU — Florida State University Herbarium, Tallahassee, Florida

GH — Gray Herbarium, Harvard University, Cambridge, Mas-

sachusetts

IJ — Institute of Jamaica, Kingston, Jamaica

K — Royal Botanic Gardens, Kew, England

MO — Missouri Botanical Garden, St. Louis, Missouri

NA — U. S. National Arboretum, Washington, D.C.

NY — New York Botanical Garden, Bronx, New York

P — Musee National d'Histoire Naturelle, Paris, France

R — Museu Nacional, Rio de Janeiro, Brazil

S-PA — Paleobotanical Department, Naturhistoriska Riksmuseum,

Stockholm, Sweden

U — Botanisch Museum, Rijksuniversiteit, Utrecht. Netherlands

US — U. S. National Museum, Washington, D.C.

YEN — Institute Botienico, Caracas, Venezuela

A number of specimens were collected in Costa Rica in 1968 on a

Field Museum trip with Dr. William Burger, the primary purpose of

which was continued general collecting for current research on the

"Flora Costaricensis."1 Some important additional collections were

1 This trip was supported by funds from National Science Foundation Grant

GB-7300, L. O. Williams, Principal Investigator.

23

24 FIELDIANA: BOTANY, VOLUME 37

made at my request, in the critical areas of Oaxaca, Mexico, by Dr. John

Mickel, and Choco, Colombia, by Dr. David Lellinger.

MAPS

Maps showing distribution of species (pages 82-85) were charted

on the Goode Series of base maps, published by the University of Chi-

cago Press, Chicago, Illinois. Each symbol represents a specific locality

where a particular species has been collected.

ACKNOWLEDGEMENTS

I wish to express my deep appreciation to Dr. Rolla Tryon for his ad-

vice and criticism throughout the course of this work, and for so un-

selfishly granting me hours of his time during my frequent visits to the

Gray Herbarium. It was, in large part, his inspiration and guidance that

provided the original impetus for this research.

My sincere thanks are also extended to Dorothy Nash for her assis-

tance in the typing and proofreading of manuscript; to Dr. Rolf Singer for

criticism and correction of the Latin descriptions; and to Dr. Thomas

Taylor (now of Ohio State University) and Mr. Ron Wibel for their

help in use of the scanning electron microscope at the University of

Illinois, Circle Campus, in Chicago, Illinois. I am indebted to T. R.

Dudley, David Lellinger, Terry Lucansky, John Mickel, Timothy Plow-

man, and Richard White for Cnemidaria collections of special impor-

tance, and to the many herbarium curators for their generous loans of

specimens used in this study. I am especially grateful to Luis D. Gomez

and other members of the staff of the Museo Nacional, San Jose, Costa

Rica, for the co-operation and courtesies extended to me while collecting

in their country.

Many specimens which I examined at Gray Herbarium had been

collected on field trips, or borrowed on visits to European herbaria, by

Drs. Rolla and Alice Tryon. These trips were supported by funds from

National Science Foundation Grants GB-4184 (Rolla Tryon) and

GB-3 1 1 70 (Rolla and Alice Tryon) .

Most of the fine drawings used as illustrations were done by Richard

Roesener of Field Museum. Others were done by Sally Babb Landry

and Lydia Vickers Wunsch at Gray Harbarium. Useful information

about the spores of many species of Cyatheaceae was obtained from

dozens of slides prepared and loaned to me by Dr. Gerald Gastony.

Finally, I wish to thank Dr. Louis O. Williams, whose advice and

encouragement have been a continual source of strength to me from

the inception of this work.

25

SYSTEMATIC TREATMENT

Cnemidaria Presl, Tent. Pterid. 56. 1836. TYPE SPECIES: Cne-

midaria speciosa Presl.

Cnemidopteris Reichenb., Deutsche Bot. 1 (Repert, Herb. Nomencl. Gen. PI.),

Abtheil. 2: 148, 235. 1841 nom. illeg., illegitimate correction of the name

Cnemidaria Presl.

Hemistegia Presl. Gefassbiindel Stipes der Farm, 46, 1847 (preprint of Abh.

Bohm Ges. Wiss. V, 5: 354. 1848). TYPE SPECIES: Hemistegia kohautiana

(Presl) Presl = Cnemidaria grandifolia (Willd.) Proctor.

Microstegnus Presl, Gefassbiindel Stipes der Farm, 45, 1847 (preprint of

Abh. Bohm Ges. Wiss. V, 5: 353, 1848). TYPE SPECIES: Microstegnus grandi-

folius (Willd.) Presl =: Cnemidaria grandifolia (Willd.) Proctor.

Actinophlebia Presl, Gefassbiindel Stipes der Farm, 47, 1847 (preprint of Abh.

Bohm Ges. Wiss. V, 5: 355, 1848). TYPE SPECIES: Actinophlebia horrida (L.)

Presl = Cnemidaria horrida (L.) Presl.

Hemitelia pro pane auctt.

Caudex ascending to erect, rudimentary to 1.5 m. long, rarely reported up to

7 cm. in diameter at base and 3.5 m. in length, bases often thickly covered with

adventitious roots; leaf to 3.5 m. long, 1.5 m. wide, broadly lanceolate to ovate-

oblong; lamina pinnate to pinnate-pinnatisect, never fully bipinnate, tissue

glabrous; petiole smooth to muricate, or provided with stout, conical spines up to

7 mm. long, with pneumatophores 0.5 to 1 cm. long, spaced 4 to 5 cm. apart in

a single line (rarely double) along either side of the petiole, spindle-shaped, linear

to lanceolate; petiole scales generally appressed, linear to ovate, or amorphous,

whitish, or more commonly bicolorous with a dark brown median stripe and a

whitish margin, marginal cells differentiated from those of the central portion also

in shape and orientation; major axes rounded beneath, sulcate above, often pro-

vided with minute, appressed, arachnoid pubescence; scales of the rachis and

minor axes scattered to rare, or lacking, broad, ovate or amorphous, whitish, pale,

brown or bicolorous, flat and appressed (or some species with bullate costal

scales); stiff, terete, recurved, pluricellular trichomes to 0.4 mm. long copious to

sparse to commonly lacking on rachis and costa, but always lacking on costa and

costules above; pinnae sessile to short-stalked, patent to ascending, or commonly

the basal pair reduced and deflexed, apices acute to acuminate, margins entire or

crenately to deeply cleft, the segments obtuse to acuminate, rarely apiculate; veins

pinnately arranged in the segments, the basal veinlet merging with the one

opposite to form a costal areole, or strongly connivent to the sinus, or (only in

C. amabilis) non-connivent and reaching the margin at a point just above the

sinus; sori subglobose at maturity, disposed on the veins in a single inframedial to

submarginal line between costule and segment margin, or in the most deeply

dissected species arranged in several lines; receptacle elevated, subglobose, with

26

STOLZE: REVISION OF CNEMIDARIA 27

paraphyses rudimentary or lacking; indusia subtending the sorus, commonly

saucer-shaped to semicircular, with margins erose-entire or several-lobed, or

rarely fully circular or reduced to one narrow lobe, or in young plants of some

species irregularly cup-shaped to subglobose; spores trilete, ca. 37-52 /j.., pro-

vided with one large pore near the center of each face on or near the equator,

and with numerous smaller apertures of irregular size and shape scattered over

the surface.

This description is an emendation of that in Tryon (1970), restrict-

ing the genus to those species with the characteristics enumerated here

and in the "Introduction." Thus the excluded species previously included

under the old generic name, Hemitelia, are herewith placed in the genus

Cyathea (See "Excluded Species").

Therefore I would emend the part of the key which includes the

genera Trichipteris, Cyathea, and Cnemidaria, in Tryon's "The Classi-

fication of Cyatheaceae," (1970) as follows:

Spores with or without apertures of various size and shape, but never

with a single very large pore near the center of each face on or

near the equator; costae and often costules with minute, terete,

recurved trichomes abundant on adaxial side; basal veins of seg-

ments commonly free (only in rare species merging to form

costal areoles), non-connivent, reaching the margin at a point

above the sinus; leaves commonly twice-pinnate or more (in rare

species once-pinnate).

Indusium absent Trichipteris

Indusium present, rarely scalelike, to sometimes hemitelioid,

to usually sphaeropteroid Cyathea

Spores always provided with one very large pore near the center of

each face on or near the equator; costae and costules lacking

trichomes on adaxial side; basal veins of segments commonly

merging to form costal areoles, or, when free, connivent to the

sinus (only in C. amabilis non-connivent); leaves pinnate to pin-

nate-pinnatisect, never fully bipinnate Cnemidaria

KEY TO THE SPECIES AND VARIETIES OF CNEMIDARIA

a. Basal veins of segments commonly free, rarely (if ever)

joining to form costal areoles. b.

b. Soral line submarginal on segments, c.

c. Segments cuspidate; pinnae mostly sessile or subsessile;

indusia narrow, usually entire. Mexico 5. C. apiculata

c. Segments rounded; pinnae often stalked; indusia

broad, semicircular to circular, with 2-3 lobes 8. C. bella

b. Soral line inframedial to supramedial between costule

and margin of segment, d.

d. Petiole scales commonly concolorous, whitish; basal

acroscopic veins of segments simple. Venezuela. ... 1. C. amabilis

d. Petiole scales commonly bicolorous; basal acroscopic

veins of segments branched, e.

28

FIELDIANA: BOTANY, VOLUME 37

e. Minute, terete, recurved trichomes abundant on

rachis beneath; segments obtuse. Panama, Costa

Rica 2b. C. mutica

var. chiricana

e. Minute, terete, recurved trichomes lacking, or when

present the segments acute to acuminate, f.

f. Terminal section of leaf nearly half the total

length of lamina; pinnae 6 pairs or less.

Colombia 4. C. singulars

f. Terminal section of leaf one-fifth the total length

of lamina (or less); pinnae on mature leaves

10 pairs or more. g.

g. Basal basiscopic veins of segments commonly

arising from costule or its junction with costa;

petiole frequently spiny. Panama, Costa Rica,

h.

h. Larger secondary segments crenate to shal-

lowly-lobed, some basal ones with broad,

rounded sinuses, or a few segments adnate. 2c. C. mutica

var. grandis

h. Secondary segments subentire (except at

serrulate tips), with acute, mostly narrow

sinuses, all segments joined, i.

pinnae more than twice as long as broad

i. Secondary segments (or lobes) of larger

pinnae more than twice as long as broad

beyond the sinus; sinuses extending more

than three-fourths to costa 2a. C. mutica

var. mutica

i. Secondary segments (or lobes) of larger

pinnae not more than \l/2 times as

long as broad beyond the sinus; sinuses

extending one-half to three-fourths to

costa 2d. C. mutica

var. contigua

g. Basal basiscopic veins of segments (especially

in basal one-third or apical one-third of

pinna) commonly arising from costa; petiole

lacking spines, j.

j. Soral line distinctly inframedial; basal pair

of pinnae not conspicuously reduced.

Colombia 3. C. tryoniana

}. Soral line medial to mostly supramedial;

basal pair of pinnae distinctly reduced, k.

k. Lamina gradually reduced to a non-con-

form, pinnatifid apex. Peru, Brasil ... 9a. C. uleana

var. uleana

k. Lamina with a conform terminal pinna.

Colombia, Ecuador 9b. C. uleana

var. abitaguensis

STOLZE: REVISION OF CNEM1DARIA 29

a. Basal veins commonly joining to form a series of costal

areoles, rarely (if ever) free. 1.

1. Pinnae (excluding the basal pair and reduced apical

ones) deeply pinnatifid, the sinuses extending more

than halfway to costa. m.

m. Rachis and petiole broadly alate throughout; large

creamy-white scales abundant along petiole. Peru. . 12. C. alatissima

m. Rachis and petiole not alate; petiole scales brown, or

bicolorous with whitish margins, n.

n. Indusia circular, completely surrounding receptacle.

Panama 10. C. cocleana

n. Indusia semicircular, positioned on costular side of

receptacle, o.

o. Lamina with a conform terminal pinna; seg-

ments of pinnae subimbricate, with sinuses

markedly cartilaginous above and contrasting

in color with adjacent tissue; soral line infra-

medial between costule and margin of segment.

Colombia, Ecuador 11. C. quitensis

o. Lamina gradually or abruptly reduced to a non-

conform, pinnatifid apex; segments with broader

sinuses, these not cartilaginous, tissue uni-

formly green; soral line medial to mostly supra-

medial, p.

p. Scales on costules beneath abundant, bullate.

lesser Antilles, northern Venezuela, q.

q. Segments mostly acute to acuminate; scales

on costae and costules beneath white to

bicolorous. St. Lucia northward to Saba. . 7a. C. grandifolia

var. .grandifolia

q. Segments mostly obtuse; scales on costae

and costules beneath brown. St. Vincent

southward to coastal Venezuela 7b. C. grandifolia

var. obtusa

p. Scales on costae and costules beneath rare or

scattered, broad, flat, never bullate. r.

r. Rachis and petiole muricate to spiny; scales

of costae and costules brown. Greater

antilles, Costa Rica, Venezuela to Peru . . 6. C. horrida

r. Rachis smooth, petiole smooth to tuber-

culate; scales of costae and costules pale

or whitish. Brazil, Peru 9a. C. uleana

var. uleana

1. Pinnae (excluding the basal pair and reduced apical

ones) entire to shallowly pinnatifid, the sinuses extend-

ing halfway or less to costa. s.

s. Pinnae (excluding the basal pair and reduced apical

ones) entire to broadly and shallowly crenate or

broadly serrate; 5-9 cm. broad. Venezuela, Peru. t.

30

FIELDIANA: BOTANY, VOLUME 37

t. Veins mostly once-forked; bases of pinnae truncate;

indusia semicircular, attached on costular side

of receptacle. Venezuela

t. Veins simple; bases of pinnae rounded to broadly

cuneate; indusia circular, completely surrounding

receptacle. Ecuador, Peru

s. Pinnae (excluding the basal pair and reduced apical

ones) deeply-crenate to shallowly pinnatifid, or if

subentire, then never over 4 cm. broad, u.

u. Lamina gradually or abruptly reduced to a pinna-

tifid apex, or with a non-conform terminal pinna,

v.

v. Pinnae (excluding the basal pair and reduced

apical ones) shallowly pinnatifid, the sinuses

extending one-third to halfway to costa; veins

commonly branched, w.

w. Narrow, membranous wing commonly extend-

ing well down each side of rachis. Costa

Rica, Colombia

w. Narrow wing lacking on the rachis below the

2 uppermost pairs of pinnae, x.

x. Lamina reduced very gradually to a pinnati-

fid apex, lacking a distinct apical section;

pinna segments often imbricate, with sin-

uses tending to be cartilaginous; pinnae

less than 4.5 cm. wide. Probably only

French Guiana

x. Lamina reduced very abruptly into a con-

spicuous, separate apical section; segments

not imbricate, or if so, the larger pinnae

5-7 cm. wide. y.

y. Indusia fully circular, the margins sub-

entire. Panama

y. Indusia more or less semicircular, the

margins erose to several-lobed. z.

z. Soral line medial between costule and

margin of segment; segments com-

monly imbricate, with sinuses cartila-

ginous and brownish colored. Colom-

bia

z. Soral line supramedial; segments usually

with broader sinuses, these rarely car-

tilaginous. Trinidad, Tobago, Mar-

garita, Guianas, Venezuela

v. Pinnae (excluding the basal pair and reduced

apical ones) subentire to deeply crenate, the

sinuses extending less than one-third to costa;

veins simple, aa.

22. C. karsteniana

23. C. nervosa

13. C. choricarpa

16. C. cruciata

10. C. cocleana

15b. C. spectabilis

var. colombiensis

15a. C. spectabilis

var. spectabilis

STOLZE: REVISION OF CNEMIDARIA 31

aa. Trichomes and scales scattered to abundant

on rachis beneath; sori inframedial on

veins. Mexico, Guatemala 21. C. decurrens

aa. Trichomes and scales lacking on rachis; sori

in a V-shaped pattern, supramedial on

veins near segment base but inframedial

near tip. British Guiana 20. C. roraimensis

u. Lamina with a conform or subconform terminal

pinna (this rarely with a pair of longer basal

lobes), bb.

bb. Soral line mostly inframedial on the veins be-

tween costule and margin of segment, cc.

cc. Petiole and often the lower rachis spiny;

minute, terete, recurved trichomes scat-

tered to abundant on rachis and costae

beneath. Colombia, Ecuador 11. C. quitensis

cc. Petiole lacking spines; axes lacking terete,

recurved trichomes. dd.

dd. Veins from the costule simple to once-

forked; scales of costa and rachis com-

monly whitish or pale; pinnae (exclud-

ing the basal pair and reduced apical

ones) mostly deeply crenate. Colombia,

Ecuador 19. C. ewanii

dd. Veins from the costule 2- or 3-forked;

scales when present commonly glossy,

dark brown; pinnae (excluding the basal

pair and reduced apical ones) pinnatifid,

the sinuses extending one-third to half-

way to costa. Colombia 14. C. chocoensis

bb. Soral line medial to supramedial between cos-

tule and margin of segment, ee.

ee. Petiole spiny or strongly muricate. Trinidad,

northern Venezuela 17. C. consimilis

ee. Petiole smooth, or rarely tuberculate at base.

Peru, Bolivia 18. C. speciosa

1. Cnemidaria amabilis (Morton) Tryon, Contr. Gray Herb. 200:

52. 1970. Figure 8b, Map 2.

Hemitelia amabilis Morton, Fieldiana Bot. 28: 10. 1951. TYPE COLLEC-

TION: Steyermark 62042, northeast of Caripe, Monagas, Venezuela, April. 13,

1945. HOLOTYPE: US! PHOTO: F! GH! ISOTYPES: F! VEN!

Caudex to 1 m. long, 3 to 7.5 cm. in diameter; leaves to 1 m. long and 0.5 m.

broad, terminating abruptly in a pinnatifid apex; petiole to 20 cm. long, with spines

to 1.5 mm. long or lacking, the scales commonly concolorous, whitish, occasionally

with a dark brown spot at the point of attachment; rachis with thin, green, her-

baceous (often inconspicuous) wing extending partly to wholly down each side,

the scales frequent, white or yellowish to light brown, or rarely bicolorous, the

32 FIELDIANA: BOTANY, VOLUME 37

trichomes minute, terete, recurved, abundant above and beneath; pinnae sessile,

cut three-fourths to costa, basal pair substantially reduced and deflexed; costae

and costules with whitish or brown scales scattered to frequent beneath, and

trichomes scattered to abundant beneath; ultimate segments subfalcate, obtuse

to rarely subacute, basal pair occasionally overlapping the rachis; veins always

free, basal ones non-connivent, and reaching the margin at a point above the

sinus, commonly once-forked (except basal acroscopic veins simple), basal

basiscopic ones arising from the costule; sori in a single line, supramedial between

costule and segment margin; indusia yellow to brown, mostly semicircular, suben-

tire to 2- or 3-lobed.

Cnemidaria amabilis is evidently the most primitive species in the

genus. Characters that are common in the other genera of scaly Cya-

theaceae — but present in Cnemidaria only in C. amabilis or a few other

primitive species — are the following: the upper side of the rachis is

closely invested with recurved, terete trichomes (also in C. mutica var.

chiricana); the pinnae are deeply dissected (ca. three-fourths of the

way to the costa); and the axes lack the peculiar arachnoid scurf

which is present on most other species of Cnemidaria. In addition the

venation is similar to that in most Cyatheaceae, not modified as in the

other species of Cnemidaria. The basal veins are completely free, ex-

tending from the costule to the margin at a point above the sinus of the

converging segments, the opposing veins having no tendency to incline

toward one another. Thus C. amabilis has fewer of the specialized char-

acters of Cnemidaria than any other species.

Within the genus, C. amabilis might be confused with C. mutica var.

chiricana of Panama and Costa Rica; however, the former has whitish

petiole scales with rarely a minute spot of brown at point of attachment,

and the basal acroscopic veins in each pinna segment are always simple

(fig. 8b), whereas the latter has bicolorous petiole scales, pale or whitish

with a broad median stripe of dark brown from base to tip, and all basal

veins of segments are once-forked.

In mountain forests, northern Venezuela, 850-1,450 m.

Additional specimens examined. — Venezuela. Sucre: Peninsula de

Paria, Cerro Humo, n.e. de Irapa, Steyermark 94886 (GH, NY, U,

VEN). Peninsula de Paria, Cerro Patao, n.e. de Giiiria, Steyermark

& Agostini 91205 (US, VEN) .

2. Cnemidaria mutica (Christ) Tryon, Contr. Gray Herb. 200: 52.

1970.

Caudex to 0.7 m. long and 6 cm. in diameter; leaves to 3 m. long and 1.5 m.

broad, terminating in a gradually pinnatifid apical section; petiole to 1.5 m. long,

with spines to 2.5 mm. long or lacking, the scales commonly bicolorous, dark

brown with a narrow, whitish margin; rachis with thin, green membranaceous

STOLZE: REVISION OF CNEMIDARIA

33

(often inconspicuous) wing extending partly to wholly down each side, or lacking

entirely, the scales scattered to frequent beneath, pale to brownish or bicolorous,

the trichomes minute, terete and recurved, abundant to lacking; pinnae sessile

to short-stalked, cut two-thirds to (rarely) entirely to the costa, basal pair

substantially reduced and often deflexed; costae and costules with pale to brown

or bicolorous scales scattered to frequent beneath, and trichomes lacking or

scattered beneath; secondary segments subfalcate to falcate, obtuse to. acuminate,

basal pair rarely overlapping the rachis; veins free, connivent to the sinus, only

rarely merging to form costal areoles, 1- to 4-forked, to pinnately branched, basal

basiscopic ones commonly arising from the costule; sori in one to several lines,

inframedial to supramedial between costule and segment margin; indusia pale

yellow to brown, more or less semicircular, subentire, erose or 2- to 3-lobed, often

irregularly cup-shaped in young material.

C. mutica has the most variable leaf outline of any species in the

genus. Of seven species treated by Maxon (1912, 1914), I can recog-

Cnemidaria mutica

FIG. 13. Variable pinna outline of Cnemidaria mutica: a, var. contigua; b-e,

var. mutica; f-h, var. grandis.

34 FIELDIANA: BOTANY, VOLUME 37

nize only one, including four varieties. The differences between these are

based primarily on segment shape and degree of dissection of the pinnae.

Although there is no difficulty in separating the two extremes (var.

contigua and var. grandis), the degree of dissections and relative shape

of the segments blend gradually and almost imperceptibly from one

variety to the next (fig. 13). Thus the varietal limits I have set are

necessarily somewhat arbitrary. The differences between members of the

C. mutica complex are mostly all quantitative, although var. chiricana

is set apart from the rest by the presence of the terete, recurved trichomes

on the rachis above. There is apparently a correlation between size of

the leaf and pinnae and the degree of dissection, shape of segment

apices, and number of forks in the veins. Thus the smaller leaf of var.

contigua has pinnae about 3 cm. broad cut three-fourths to the costa,

with rounded segments, and veins only once- to twice-forked, whereas

the much larger leaf of var. grandis has pinnae often up to 15 cm. broad,

sometimes cut entirely to the costa, with mostly acuminate secondary

segments, and veins several-forked to pinnately branched. The lack

of costal areoles and the deep dissection of pinnae indicate that this

species is among the most primitive in the genus. In var. chiricana, the

presence of trichomes on the rachis above indicates that it is closely

related to the Venezuelan C. amabilis, which it also resembles in numer-

ous other features. All varieties of C. mutica are found growing at

middle elevations, encompassing only a narrow geographic range: four

Costa Rican provinces, and the neighboring Chiriqui area of Panama.

It is presumed that these are incipient species, but that reproductive

isolation has been insufficient to result in morphological distinction.

2a. Cnemidaria mutica var. mutica. Figure 7b, Map 4.

Hemitelia mutica Christ, Bull. Soc. Bot. Geneve II. 1: 233. 1909. TYPE COL-

LECTION: Werckle s.n., Turrialba, Costa Rica, 850 m. FRAGMENT: US!

Hemitelia arachnoidea Maxon, Contr. U.S. Natl. Herb. 16: 34. 1912. TYPE

COLLECTION: Maxon 453, vicinity of La Palma, Costa Rica, May 6 to 8,

1906. HOLOTYPE: (2 sheets) US! ISOTYPE: NY!

Hemitelia horrida (L.) R.Br. var. heterosora Rosenst., Repert. Spec. Nov.

Regni Veg. 10: 275. 1912. TYPE COLLECTION: Brade 451 luan Vinas prope

Rio Chis, 20/111/1910. FRAGMENT: US!

Hemitelia pittieri Maxon, Contr. U.S. Natl. Herb. 16: 32. 1912. TYPE COL-

LECTION: Pittier 10969 pro parte, Canas Gordas, Valle de Agua Buena, Costa

Rica, Feb. 1897. HOLOTYPE: US! FRAGMENT: F!

Hemitelia snbglabra Maxon lorn. cit. 36. TYPE COLLECTION: Maxon 451,

vicinity of La Palma, Costa Rica, May 6 to 8, 1906. HOLOTYPE: US!

Cyathea mutica (Christ) Domin, Pteridophyta 264. 1929.

Cyathea pittieri (Maxon) Domin, loc. cit.

STOLZE: REVISION OF CNEMIDARIA 35

Cyathea snbarachnoidea Domin, loc. cit., nom.nov. for Hemitelia arachnoidea

Maxon.

Cyathea subglabra (Maxon) Domin, loc. cit.

Petiole spines to 2.5 mm. or, rarely, lacking; rachis commonly lacking the

minute, terete, recurved trichomes; larger pinnae cut more than three-fourths

to costa; scondary segments subentire (except at serrulate tips), obtuse to

acuminate, joined by narrow, acute sinuses, the segments of larger pinnae more

than twice as long as broad beyond the sinus; veins 1- to 4-forked; sori in 1 or

2 series between margin of segment and costule.

In addition to the specimen of Pittier 10969, listed above as the holo-

type of Hemitelia pittieri, there is another at Museo Nacional in San

Jose, Costa Rica labeled Pittier 10969, also identified as H. pittieri. The

latter is evidently a mixed collection. With its pinnae incised only one-

third of the way to the costa, and its abundant costal areoles, it is obvi-

ously Cnemidaria choricarpa. Maxon's description (1912) of H. pittieri

was based solely on the single sheet he saw at U.S. National Herbarium,

so he was apparently unaware of the existence of the other similarly

labeled collection in San Jose. Since Pittier 's No. 10966, also collected

at Cafias Gordas, is C. choricarpa, it may be assumed that one of the

leaves of this plant was somehow mixed with the other collection and

therefore received the number 10969 in error.

Larger specimens of var. mutica may resemble Cnemidaria horrida;

comparisons are presented under the discussion of C. mutica var. grandis.

Forests and edges of forests, 450-2,100 m., Costa Rica (Alajuela,

Cartago, Heredia, San Jose) and Panama (Chiriqui)-

Selected specimens examined. — Costa Rica. Alajuela: Along Rio San

Rafael, Aguas Zarcas, L. O. Williams et al. 29109 (F). Cartago: La

Estrella, Standley 39349 (GH, US) . Finca El Muneco, South of Cartago,

Stork 4756 (GH, US). South of Tapantf above Rio Grande de Orosi,

Burger & Stolze 5685 (F, GH). Heredia: Vara Blanca de Sarapiqui,

Skutch 3262 (GH, US). Roadside waterfall near Vara Blanca, Burger

& Stolze 5936 (F, GH). Cinchona, above Upper Sarapiqui Valley,

Scamman 7575 (GH, US). San Jose: La Palma, northeast of San Jeron-

imo, Burger & Stolze 5328 (F, GH). Lisiere des paturages a La Palma,

Tonduz 12532 (CR, US). Panama. Chiriqui: 5 miles east of El Boquete,

Killip 5156 (GH, MO, US). Vicinity of El Boquete, Common 1285

(US).

2b. Cnemidaria mutica var. chiricana (Maxon) Stolze, stat. et comb,

nov. Figure 8a, Map 5.

Hemitelia chiricana Maxon, Contr. U.S. Natl. Herb. 16: 33. 1912. TYPE COL-

LECTION: Maxon 5519 (as "H. chiriquana"), between Alto de la Palmas and

36 FIELDIANA: BOTANY, VOLUME 37

top of Cerro de la Horqueta, Chiriqui, Panama, March 18, 1911. HOLOTYPE: (3

sheets) US!

Cyathea chiricana (Maxon) Domin, Pteridophyta, 263. 1929.

Petiole spines lacking; rachis with minute, terete, recurved trichomes abun-

dant beneath, scattered to abundant above; pinnae cut two-thirds to four-fifths to

costa; secondary segments subentire (except at serrulate tips), obtuse to rarely

subacute, joined by narrow, acute sinuses, segments of larger pinnae \lA-2 times

as long as broad beyond the sinus; veins 1- to 2-forked; sori in single line be-

tween margin of segment and costule.

Var. chiricana may be easily confused with C. amabilis; the two are

compared in the discussion of the latter species.

Forests and wooded slopes, 1,400-2,200 m., Costa Rica (Cartago)

and Panama (Chiriqui) .

Additional specimens examined. — Costa Rica. Cartago: Rio Grande

de Orosi beyond Tapanti, White & Lucansky 196840 (GH). Panama.

Chiriqui: Near the divide, above El Boquete, Killip 5347 & 5350 (US).

Cordillera above El Boquete, Killip 5297 (GH, US), and Killip 5294

(US). Between Alto de las Palmas and Cerro de la Horqueta, Maxon

5474 & 5521 (US) and Maxon 5517 (GH, US) .

2c. Cnemidaria mutica var. grandis (Maxon) Stolze, stat. et comb,

nov. Figure 13, Map 6.

Hemitelia grandis Maxon, Contr. U.S. Natl. Herb. 16: 37. 1912. TYPE COL-

LECTION: Maxon 307, vicinity of Coliblanco, Costa Rica, April 30-May 2, 1906.

HOLOTYPE: (2 sheets) US! ISOTYPE: US!

Hemitelia rudis Maxon, op. cit. 17: 413, 1914. TYPE COLLECTION: Maxon

5682, Holcomb's Trail, above El Boquete, Chiriqui, Panama, March 23, 1911.

HOLOTYPE: (4 sheets) US! ISOTYPE: GH!

Cyathea grandis (Maxon) Domin, Pteridophyta, 264. 1929.

Cyathea rudis (Maxon) Domin, loc.cit.

Petiole spines to 2.5 mm. or, rarely, lacking; rachis with minute, terete, re-

curved trichomes lacking above, scattered to lacking beneath; pinnae cut nearly

to costa (or larger ones fully to costa at base); larger secondary segments cre-

nate to shallowly-lobed, some basal ones joined by broad, rounded sinuses, or a

few segments adnate, most segments twice to four times as long as broad beyond

the sinus; veins thrice-forked to pinnately branched; sori in 1-4 series between

margin of segment and costule.

In general size and pinna architecture, leaves of var. grandis and

larger leaves of var. mutica closely resemble those of C. horrida and C.

grandifolia var. grandifolia. Also, although the former two are basically

free- veined and the latter two commonly have basal veins merging to

form costal areoles, occasional specimens have variation in these char-

STOLZE: REVISION OF CNEMIDARIA 37

acters. For example, rarely a pinna of C. mutica may have a number of

costal areoles, or occasionally a pinna of C. horrida or C. grandifolia

may have segments with a number of free basal veins. However, in

C. mutica (except in var. contigua) the soral line or lines are roughly

equidistant between margin and costule, or, in the largest specimens,

nearly fill both sides of the secondary segment. The soral lines of C.

horrida or C. grandifolia extend to very near the margin, a rather dis-

tinctive character shared within the genus only by C. bella and C. apicu-

lata. Cnemidaria grandifolia var. grandifolia is further distinguished by

the costules which bear abundant white, bullate scales often terminating

in a hairlike tip, a feature peculiar to this species.

Forest and wooded slopes, 1,250-1,900 m., Costa Rica (Cartage, San

Jose) and Panama (Chiriqui).

Selected specimens examined. — Costa Rica. Cartago: Valley of

Rio Grande de Orosi, Tryon & Tryon 7036 (GH); Near Tapanti, Lent

28 (F, GH). El Muneco, on the Rio Navarro, Standley & Torres 51250

(F, US). El Cedral, near Naranjo River, Nisman 123 (GH). South

slope of Turrialba Volcano, L. O. Williams 19647 (US). San Jose: La

Palma area, northeast of San Jeronimo, Burger & Stolze 5270 (F, GH).

Vicinity of La Palma, Maxon & Harvey 8023 & 8042 (US). Panama.

Chiriqui: 10 miles above El Boquete, Killip 5231 & 5248 (US). Valley

of Rio Piarnasta, above El Boquete, Killip 5391 (GH, MO, US) .

2d. Cnemidaria mutica var. contigua (Maxon) Stolze stat. et comb,

nov. Underw. ex Maxon Figure 13, Map 7.

Hemitelia contigua Maxon, Contr. U.S. Natl. Herb. 16: 32. 1912. TYPE COL-

LECTION: Maxon 523, 5 mi. south of Cartago, Costa Rica, May 12, 1906.

HOLOTYPE: (2 sheets) NY! FRAGMENT: US!

Cyathea contigua (Maxon) Domin, Pteridophyta 263. 1929.

Petiole spines minute or commonly lacking; rachis lacking minute, terete, re-

curved trichomes; pinnae cut one-half to three-fourths to costa; secondary segments

subentire (except at serrulate tips), obtuse to rarely subacute, joined by narrow,

acute sinuses, the segments not more than \1A times as long as broad beyond the

sinus; veins 1- to 2-forked; sori in a single line between margin of segment and

costule.

In addition to the differences noted in the key, var. contigua may also

be distinguished by the V-shaped soral pattern on the segments (the

apical sori situated very near the costule, while the basal ones are quite

close to the margin), whereas the sori in other varieties of C. mutica

are relatively constant in their position between margin and costule,

whether inframedial, medial, or supramedial. Trichomes are completely

lacking in var. contigua, and scales on all axes except the petiole are

38 FIELDIANA: BOTANY, VOLUME 37

uniformly brown, whereas scales may be pale to brown, or commonly

bicolorous, in other varieties of C. mutica.

Mountain forests, 1,250-2,100 m., Costa Rica (Cartago, Heredia).

Additional specimens examined. — Costa Rica. Cartago: Santa Clara

de Cartago, Maxon & Harvey 8220 (US). La Sierra, 25 km. south of

Cartago, L. O. Williams et al. 28016 (CR, F). Heredia: Socorro de

San Ramon, Brenes 4992 (F). Forests of Rio Vueltas, Gomez 2246

(CR, F, GH, NY, US). Slope of Volcan Barba, Scamman & Holdridge

7879 (GH, US).

3. Cnemidaria tryoniana Stolze, sp. nov. Figures lOc, 14; Map 2.

Caudex brevissimus vel nullus; petiolus inermis; pinnae infimae vix deminutae

vel deflexae; costae subtus paleis latis niveis adpressis instructae; venae semel

furcatae, liberae; venae infimae baud anastomosantes sed arcuatae et in sinum

conniventes; venae infimae basioscopicae e costa egredientes; sori uniseriati, inter

costulam et marginem inframediani.

TYPE COLLECTION: Metcalf & Cuatrecasas 30122, between Valdivia and

Yarumal, Antioquia, Colombia, Feb. 20, 1942. HOLOTYPE: (2 sheets) US!

PHOTO: F! ISOTYPES: GH! MO! FRAGMENT: F!

Caudex very short or lacking; leaves to 1.8 m. long and 0.5 m. broad, termi-

nating abruptly with a subconform apical pinna; petiole to 0.7 m. long, unarmed,

the scales sparsely scattered, lanceolate, bicolorous, dark shiny brown with nar-

row, whitish margins; rachis non-alate, the scales scattered, broad, dirty white

or occasionally with dark brown centers, trichomes lacking; pinnae distant, sessile,

cut three-fourths to costa, basal pair scarcely (if at all) reduced or deflexed;

costae and costules with whitish scales scattered to frequent beneath, trichomes

lacking; ultimate segments subfalcate, subacute to acute, basal pair commonly

crowding or overlapping the rachis; veins free, once-forked, basal ones conniving

at the sinus, basal basiscopic ones arising from the costa; sori in a single line,

inframedial between costule and segment margin; indusia yellow to brownish,

mostly semicircular and subentire.

This species is one of the most primitive in the genus, as evidenced

by its free venation and deeply dissected pinnae, and is closely related

to C. amabilis and C. singularis. It could perhaps be confused with

C. uleana (especially var. abitaguensis) , which also has unarmed petioles

and basal veins which arise from the costae rather than from the costules.

However, in addition to characters noted in the key, C. tryoniana can

further be distinguished by the abundance of large, whitish scales along

the costae beneath, whereas the scales of C. uleana are rare or sparse.

It is with great personal satisfaction that I name this species in honor

of Dr. Rolla M. Tryon, Jr., who has done excellent work with the

ferns of tropical America, and who is currently involved in a comprehen-

sive study of the Cyatheaceae.

FIG. 14. Cnemidaria tryoniana, Metcalf & Cuatrecasas 30122 (US, holotype).

39

40 FIELDIANA: BOTANY, VOLUME 37

The species is known only from the type collection; found at an al-

titude of 2,200 m.

4. Cnemidaria singular!* Stolze, sp. nov. Figure 15, Map 8.

Caudex brevissimus vel nullus; petiolus inermis; folii sectio terminalis dimidiae

parti longitudinis laminae aequalis; pinnae 3-6-jugatae; costae subtus paleis latis

niveis adpressis sparsis instructae; venae furcatae vel bifurcatae, liberae; venae

infimae baud anastomosantes sed arcuatae et in sinum conniventes; venae infimae

basioscopicae e costa egredientes; sori uniseriati, inter costulam et marginem

medii vel supramediani.

TYPE COLLECTION: Soejarto 1571, Cerro Portachuelo, Depto. de Putu-

mayo, Colombia, Aug. 27, 1965. HOLOTYPE: GH! PHOTO: F! ISOTYPE: F!,

GH!

Caudex very short or lacking; leaves to 1.2 m. long and 0.4 m. broad, pinnati-

fid apical section nearly half the total length of the lamina; petiole to 0.6 m.

long, unarmed, the scales sparse except at very base, lanceolate to linear-lanceo-

late, bicolorous, dark shiny-brown with narrow whitish margins; rachis non-alate,

the scales rare, thin, very pale brown or whitish, trichomes lacking; pinnae

distant, six pairs or less, sessile, commonly cut a little more than halfway to

costa, basal pair slightly reduced and somewhat de flexed; costae and costules

with thin, whitish, amorphous scales rare to scattered beneath, trichomes lacking;

ultimate segments subfalcate, rounded to subapiculate; veins free, once- or twice-

forked, basal ones conniving at sinus, basal basiscopic ones arising from the

costa; sori in a single line, medial to supramedial between costule and segment

margin; indusia yellowish-brown, mostly semicircular, subentire to deeply two-

lobed.

This shares with C. tryoniana the more primitive characteristics of

the genus. The two species also have in common a number of other

morphological features, notably: the rudimentary caudex, the lack of

spines of the petiole and trichomes on the axes, the basal basiscopic veins

arising from the costa rather than from the costules, and the scales on

the axes (when present) thin, translucent, and whitish. However, C.

singularis is one of the most unique species in the genus, and cannot be

confused with others. Its laminar architecture is almost monstrous in

that the terminal section measures nearly half the length of the entire

lamina, while the remainder is comprised of only 3-6 pairs of pinnae.

The leaves therefore present a strange, top-heavy aspect.

The species is known only from the type collection, which was found

at 2,300 m. in a wet mountain forest, along the road from Sibundoy to

Pepino, Dept. Putumayo, Colombia.

5. Cnemidaria apiculata (Hook.) Stolze, comb. nov. Map 3.

Hemitelia apiculata Hook. & Bak., Syn. Fil. 29, 1865. TYPE COLLECTION:

Jurgensen 873, Sierra San Pedro Nolasco, Talea, etc., Oaxaca, Mexico, 1843-

PLANTS OF COLOMBIA

FIELD MUSEUM OF

NATURAL HISTORY

FIG. 15. Cnemidaria singulars, Soejarto 1571, (GH, holotype).

41

42 FIELDIANA: BOTANY, VOLUME 37

1844. HOLOTYPE: K, FRAGMENTS: NY! US! ISOTYPE: P! PHOTO: F!

Cyathea aristata Domin, Acta Bot. Bohem. 9: 93. 1930, nom. nov. for Hemi-

telia apiculata Hook., (non Cyathea apiculata Domin, 1929).

Caudex not seen; leaves to 1.5 m. long and 0.7 m. broad, terminating in a

gradually pinnatifid apical section; petiole to 0.7 m. long, spines lacking or, very

rarely, minute, scales frequent, bicolorous, dark brown with narrow whitish

margins, trichomes lacking; rachis non-alate, the scales scattered to abundant,

bicolorous, trichomes lacking; pinnae rather crowded, sessile, cut three-fourths

to seven-eighths to costa; costae and costules with minute, amorphous, brown or

bicolorous scales scattered to lacking beneath, trichomes lacking; ultimate seg-

ments falcate, sharply cuspidate, basal pair commonly overlapping the rachis;

veins free, basal ones conniving at the sinus, basal basiscopic ones arising from

the costule; sori in a single line, submarginal on the segments; indusia brown,

subentire, narrow, spathulate or cucullate.

Cnemidaria apiculata is the northernmost representative of the

species related to C. amabilis. It is probably derived from the same an-

cestral stock as the Central American C. mutica, which it resembles in

a number of characters. However, C. apiculata is unique in two respects:

no other species has such sharply cuspidate segments nor such narrow

indusia (fig. 11 a). It is further distinguished by having a soral line so

far from the costule as to usually appear almost marginal (fig. lOa). This

characteristic occurs elsewhere only in C. bella, C. grandifolia, and C.

horrida. In the original description of Hooker the number "273" is given

for Jurgensen's collection. This was probably a printer's error, for the

isotype and fragments I have seen are all clearly marked "Jurgensen

No. 873", and Maxon (1912) gives this same number for the holotype

at Kew.

In rain forests, 1,200-1,600 m., found only in the State of Oaxaca,

Mexico.

Additional specimens examined. — Mexico. Oaxaca: Ridge between

Yetzelalag & Lovani, Depto. Choapam, Hallberg 1557 (NY) ca. 20

miles n.e. of Villa Alta, Distr. of Villa Alta, Michel 994 (NY) 79 km.

west of Ixtlan de Juarez on Rt. 175, Depto. Ixtlan, Michel 5676, 5717

(NY) 24 km. south of Valle Nacional, Depto. Tuxtepec, Michel 5935

(NY).

6. Cnemidaria horrida (L.) Presl, Tent. Pterid. 57. 1836. Map 10.

Polypodium horridum L., Sp. PI. 1092. 1753. TYPE: Plumier Descr. PI. Amer.

1693, t.4 (Fil. Amer. t.S. 1705).

Cyathea horrida (L.) I.E. Sm., Mem. Acad. Roy. Sci. (Turin) 5: 416, 1793.

(nonKaulf. 1823).

STOLZE: REVISION OF CNEMIDARIA 43

Cyathea commutata Sprengel, Anleit. Kennt. Gewasche 3: 146, fig. 32, 1804.

TYPE COLLECTION: Santo Domingo.

Hemitelia horrida (L.) R.Br., Prod. Fl. Nov. Holl. 158, 1810. (non H.

horrida var. imrayana Hook, in Hook. & Bak. 1865.)

Actinophlebia horrida (L.) Presl, Gefassb. Stipes der Farm 48: 1847. (pre-

print of Abh. Bohm Ges. Wiss V. 5: 356. 1848.)

Hemitelia hookeri Presl, torn. cit. 350. nom. mid.

Hemistegia horrida (L.) Fee, Mem. Fam. Foug. 5 (Gen. Fil.): 351. 1850-52.

Hemistegia repanda Fee, loc. cit. TYPE COLLECTION: Linden, s.n., Cuba.

Cormophyllnm horridnm (L.) Newm., Phytologist 5: 238. 1854.

Hemitelia acuminata Schlecht., Bot. Zeit. 14: 474, 1856. nom. nud.

Hemitelia commutata (Sprengel) Schlecht., loc. cit., nom. nud.

Hemitelia hookeriana Schlecht., loc. cit.. nom. nud.

Caudex to 4 m. tall and 7 cm. in diameter; leaves to 3.5 m. long and 1.5 m.

broad, terminating gradually in a nonconform, pinnatifid apex; petiole to 1.5 m.

long, muricate or, more commonly, with stout spines to 5 mm. long, the scales

lanceolate, bicolorous, dark brown with narrow whitish margins, usually found

only at base; rachis non-alate, often spiny, the scales rare, when present lanceolate

to ovate, dark brown or bicolorous, trichomes lacking; pinnae sessile to short-

stalked, cut seven-eighths to (rarely) entirely to the costa, basal ones slightly re-

duced and deflexed; costae and costules with dull brown, amorphous scales rare

to scattered beneath, trichomes lacking; secondary segments subfalcate or falcate,

acute to acuminate, basal pair often overlapping the rachis; veins 1- to 5-forked,

basal ones commonly merging to form costal areoles, basal basiscopic ones usually

arising from the costule; sori in one to several lines, supramedial to submarginal

between costule and segment margin; indusia whitish to yellow, more or less

semicircular, subentire to erose, saucer-shaped or sometimes, in young material,

irregularly cup-shaped to subglobose.

Because of its huge size and deeply-dissected pinnae, Cnemidaria

horrida is perhaps more representative of the typical aspect of tree ferns

than any other species of the genus. It is evidently the most primitive

of the areolate-veined group, and from its ancestral stock was probably

derived not only the other Antillean species, C. grandifolia, but most

of the South American areolate species as well.

Cnemidaria horrida is geographically the most successful species in

the genus, enjoying the widest distribution as well as the greatest al-

titudinal range. It is common in the Greater Antilles, and ranges along

the subcoastal regions of northern South America southward down the

Andes to southern Peru, and has been found at times in Costa Rica.

Larger specimens of the typical variety of Cnemidaria grandifolia var.

grandifolia, especially those with acuminate segments, may be confused

with C. horrida. Besides the scale differences noted in the key, however,

the latter has whitish or pale yellow indusia and the lamina beneath has

some white, arachnoid, scurfy indument. The indusia of C. grandifolia

44 FIELDIANA: BOTANY, VOLUME 37

var. grandifolia are most often dark- or yellowish-brown, and the lamina

lacks arachnoid scurf.

Some specimens of C. mutica in Costa Rica exhibit enough costal

areoles to be confused with C. horrida. However, the leaf texture of

C. horrida is quite thin, and the lines of sori crowd the margin closely,

whereas the leaves of C. mutica are much thicker, and the soral line

medial to supramedial (never closely approaching the margin). I have

found only two collections of C. horrida from Costa Rica; several pre-

vious determinations were of C. mutica. Maxon (1912) cities another

Costa Rican collection (Alfaro 108) as C. horrida, but I have not seen

this.

In and at the edges of forests, along stream banks and on mountain-

sides, from sea level to 2,000 m., Greater Antilles, Costa Rica, and

Venezuela to Peru.

Selected specimens examined. — Puerto Rico: Alto de La Bandera,

near Adjuntas, N. L. Britton & Shafer 2075 (F, MO, NY, US).

Lares, in sylva ad Buenos Aires, Sintenis 6088 (F, GH, NY, US).

Dominican Republic. Barahona: Vicinity of Paradis, Abbott 1618 (GH,

NY, US). Santiago: San Jose de las Matas, Valeur 350 (F, MO, NY,

US). Santo Domingo: Vicinity of Colonia Ramfis, Allard 14303 (US).

Haiti: Massif du Nord, Morne Brigand, Ekman H-2875 (F, US). Vicin-

ity of Port de Paix, E. C. Leonard 12275 (GH, NY, US). Cuba: San-

tiago de Cuba, Linden 1738 (F, US). Las Villas: Buenos Aires, Trini-

dad Mountains, Morton 4240 (GH, US). Oriente: Monteverde, Yateras,

Maxon 4336 (GH, NY, US). Pinar Del Rio: Sierra cerca de Taco Taco,

Baker 3835 (F, NY, US). Jamaica. Portland: Valley of Trafalgar River,

Maxon & Killip 779 (F, GH, NY, US). St. Andrew: Near Heritage Dam,

Proctor 3911 (US). St. Thomas: Between House Hill and Cuna Cuna

Gap, Maxon 8879 (F, GH, NY, US). Costa Rica. Heredia: Pies de Sta.

Barbara, Pittier 1679. Along Rio Sarapiqui, between Cariblanco and

San Miguel, L. O. Williams 20304 (F, US). Venezuela. Aragua: Selvas

tropofilas de Guamitas, Parque Nacional, Delgado 81 (F, US, VEN).

Distrito Federal: Cortada del Guayabo, Tamayo 154 (US, VEN). Lara:

En selvas nubladas, San Isidro, Tamayo 2543 (US, VEN). Miranda:

Near Agua Fria, km. 26, road to Carascua, Pittier 11510 (US, VEN).

Colombia. Antioquia: Porcecito, Bro. Daniel 913 (US). Caldas: San

Bernardino, Cordillera Central, Killip & Hazen 10159 (GH, NY, US).

Caqueta: Sucre, Juzepczuk 6510 (US). Cundinamarca: Arriba de Sa-

saima, Dugand & Jaramillo 3813 (US). Magdalena: Dibulla, Seifriz

277 'x (US). Meta: Along Rio Guatiquia, Villaviciencio, Pennell 1577

(F, GH, MO, NY, US). Putumayo: Between Umbria & Urcusique,

STOLZE: REVISION OF CNEMIDARIA 45

Ewan 16790 (GH, US). Santander: Jordan, 10 km. S.E. of Landazuri,

Ewan 15669 (GH). Ecuador. Napo-Pastaza: Tena, Asplund 9230 (US).

Cerro Antisana, trail to Tena, Grubb et al. 1392 (NY, US). Near Archi-

dona, Mexia 7320 (F, GH, US). Santiago-Zamora: Primary rain forest,

Taisha, Cazalet & Pennington 7711 (NY, US). Peru. Amazonas: Valley

of Rio Maranon, Bagua, Wurdack 1991 (US). Cuzco: Kosnipata-Pilco-

pata, Prov. Paucartambo, Vargas 11269 (GH). Loreto: Pumayacu, be-

tween Balsapuerto & Moyobamba, Klug 3182 (F, GH, MO, NY, US).

San Martin: San Roque, LI. Williams 7156 (F, NY, US).

7. Cnemidaria grandifolia (Willd.) Proctor, Rhodora 63: 31. 1961.

Caudex to 1 m. long and 4 cm. in diameter; leaves to 3 m. long and 1 m. broad,

terminating in a gradually- or abruptly-reduced pinnatifid apical section; petiole to

1.3 m. long, muricate, or with spines to 3 mm. long, the scales bicolorous, dark

to light brown with a whitish margin, linear to lanceolate, commonly with long

attenuate tips; rachis non-alate, with scales scattered to abundant, ovate to lanceo-

late, brown or white or bicolorous, the minute, terete, recurved trichomes scattered

to abundant beneath, or lacking; pinnae sessile, cut three-fourths to nine-tenths

to the costa, basal pair usually somewhat reduced and deflexed; costae with

broad brown, white or bicolorous scales beneath, costules with white or brown

bullate scales beneath, trichomes rare or lacking; secondary segments falcate,

obtuse to acuminate, basal pair often overlapping the rachis; veins 1- to 4- (6)

times forked, basal ones commonly merging to form costal areoles, basal basi-

scopic ones usually arising from the costule; sori in one or two lines, supramedial

to submarginal between costule and segment margin; indusia yellow-brown to

dark brown, semicircular, s-ubentire to several-lobed.

Cnemidaria grandifolia has traditionally been divided into three

species, Hemitelia grandifolia, H. obtusa, and H. kohautiana, based

mainly on the quantitative differences of pinna width and (or) shape of

the secondary segments. However, comparison of a large series of

collections will demonstrate that such characters blend almost insensibly

throughout the species and hence cannot provide a basis for taxonomic

delimitation. Pinna width varies from less than 3 cm. to more than 10

cm., while the segment apices vary from broadly rounded to long-

acuminate; and in every other morphological character examined I have

found no further evidence to warrant distinction at the species level.

"Hemitelia kohautiana," including the type collection and most speci-

mens so determined in herbaria, shares geographic distribution and

morphological characters with H. grandifolia. However, the plants

previously treated as Hemitelia obtusa differ sufficiently from the typical

Cnemidaria grandifolia to warrant varietal distinction. The pinnae are

generally much narrower in var. obtusa, and the segments are primarily

obtuse, although enough specimens can be found with wider pinnae and

46 FIELDIANA: BOTANY, VOLUME 37

at least subacute segments to be confused with similar specimens

in the typical variety. In this case, color of the bullate scales along the

costules beneath affords a more effective varietal delimitation: those of

var. obtusa are glossy and deep amber to castaneous, while those of var.

grandifolia are dull and white. Thus the combination of scale and seg-

ment characters affords an adequate basis for recognition of the two

varieties, and these are correlated with an interesting disjunction in the

Lesser Antilles. Variety grandifolia is found from Saba southward to

St. Lucia, while the range of var. obtusa begins at St. Vincent, the next

island south, and continues on to the Island of Margarita and the Para-

guana Peninsula in Venezuela. Hence there is a clear disjunction be-

tween the varieties at St. Vincent Passage. This is no greater water barrier

than others which occur among the Lesser Antilles, so the isolation of

the two taxa within C. grandifolia presumably has an environmental

basis, possibly related to the more recent emergence of the southernmost

islands after the Pleistocene.

7a. Cnemidaria grandifolia var. grandifolia. Figure 7a, Map 10.

Cyathea grandifolia Willd., Sp.Pl. 5: 490. 1810. TYPE COLLECTION: Herb.

Willd. No. 20167, Habitat in America Calidiore. HOLOTYPE: B; PHOTO: F!

GH!; FRAGMENT: US!

Hemitelia grandifolia (Willd.) Sprengel, Syst. Veg. 4: 125. 1827.

Cnemidaria kohautiana Presl, Tent. Pterid. 57. 1836. TYPE COLLECTION:

Sieber (Flora Martin. 375), Martinique; HOLOTYPE: PR or PRC? ISOTYPES:

MO! P!; FRAGMENT: US!

Hemitelia monilifera J.Sm., London J. Bot. 1: 662. 1842. nom.nud.

Hemitelia serrata J.Sm., loc.cit. nom.nud.

Hemitelia imrayana Hook., Icon. PI. VII, t.669. 1844 (non Cyathea imrayana

Hk. 1844). TYPE COLLECTION: Imray 14, Dominica; ISOTYPE: P!

Hemitelia kohautiana (Presl) Kunze, Bot. Zeit. 2: 298. 1844.

Hemistegia grandifolia Presl, Gefassb. Stipes der Farm, 47. 1847 (preprint

of Abh. Bohm Ges. Wiss. V. 5: 355. 1848). TYPE COLLECTION: Plumier tab.

26. (non Hemitelia grandifolia (Willd.) Sprengel).

Microstegnus grandifolius (Willd.) Presl., tom.cit. 46.

Hemistegia kohautiana (Presl) Presl., tom.cit. 47.

Hemistegia willdenowii Fee, Mem. Fam. Foug. 5 (Gen.Fil.): 351. 1850-52

nom.nov. for Cyathea grandifolia Willd.

Hemitelia horrida var. imrayana Hook., in Hook. & Bak., Syn. Fil. 28, 1865.

Hemistegia insignis Fee, Mem. Fam. Foug. 11: 99, 1866 (non Cyathea insignis

Eat. 1860). TYPE COLLECTION: Herb. 1'Herminier s.n., Flore de la Guade-

loupe, ISOTYPE: P!

Hemitelia insignis (Fee) C.Chr., Index Fil. 349, 1905.

Cyathea kohautiana (Presl) Domin, Pteridophyta 264. 1929.

Hemitelia obtusa var. kohautiana (Presl) Domin, Mem. Roy. Czech. Soc. Sci.

2: 71. 1929.

STOLZE: REVISION OF CNEMIDARIA 47

Cyathea antillana Domin, Acta. Bot. Bohem. 9: 91. 1930. nom.nov. for

Hemitelia imrayana Hk.

Cyathea obtusa var. kohautiana (Presl) Domin, tom.cit. 142.

Leaves terminating in a gradually pinnatifid apical section; petioles muricate

to short-spiny; rachis and costae with white to bicolorous scales; pinnae cut four-

fifths to nine-tenths to the costa; costnles with dull white, bullate scales occasional

to abundant beneath; secondary segments falcate, mostly subacute to acuminate

(only rarely obtuse); veins 1- to 4- (6-) times forked; .von in 1 to 2 lines be-

tween costule and segment margin.

In and at edges of forests on stream banks and mountainsides: 300-

1,100 m. Saba, St. Kitts, Nevis, Montserrat, Guadeloupe, Dominica,

Martinique, St. Lucia.

Selected specimens examined. — Saba: Mountain, 800 m., Boldingh

2221 ( == 7427) (IJ, U). Summit of The Mountain, Staffers 4203, (U,

US). St. Kitts: Slopes of Mount Misery, TV. L. Britton & Cowell 510

(NY, US). N.W. rim of The Crater, Proctor 19501 (GH, IJ). Nevis:

Summit of Nevis Peak, Proctor 19318 (GH, IJ). Nevis Peak, A. C.

Smith 10516 (IJ, US). Montserrat: Chaner's Mountain, Shafer 283 (F,

NY, US). Guadeloupe: Bois du Haut Matouba, Duss 4449 (F, GH, MO,

NY, US). Dominica: Sylvania Estate, alt. 488 m., Hodge 10 (GH, NY,

US). Parish of St. Paul, Wilbur et al. 8185 (F. NY). Martinique: Duss

4435 (F, GH, MO, NY, US) and Duss 1607 (NY, US). Camp de

Colson, M. & H. Stehle 4605 (US) St. Lucia: Milette Bridge, Box 490

(US). Upper slopes of Piton Flore, Proctor 18092 (GH, IJ).

7b. Cnemidaria grandifolia var. obtusa (Kaulf.) Stolze, stat. et comb,

nov. Map 10.

Hemitelia obtusa Kaulf., Enum. Fil. 252, 1824. TYPE COLLECTION: "Habitat

in Antillis." POSSIBLE SYNTYPES: Ryan s.n., Montserrat; Sieber //. mixta 331.

Cyathea mitnita Kaulf., tom.cit. 260. nom.nud.

Cnemidaria munita Presl. Tent. Pterid. 57. 1836. nom.nnd.

Cnemidaria obtusa (Kaulf.) Presl, I.e.

Hemitelia munita Hk., Sp. Fil. 1 : 32. 1844. nom.nud.

Hemistegia munita Presl, Gefassb. Stipes der Farm 47. 1847. (Preprint of Abh.

Bohm. Ges. Wiss. V. 5: 355. 1848.) nom.nud.

Hemistegia obtusa (Kaulf.) Presl. I.e.

Hemitelia munita Kuhn, Linnaea 36: 162. 1869. TYPE COLLECTION: Willd.

Herb. No. 20168, America. ISOTYPE: US!

Hemitelia bullata Christ, Bot. Jahrb. Syst. 24: 81. 1897. TYPE COLLECTION:

Eggers 6035, Grenada, 1400'. 28/XI/1889. ISOTYPES: F! P! US! FRAGMENT:

NY!

Hemitelia obtusa var. bullata (Christ) Domin, Mem. Roy. Czech. Soc. Sci. 2:

71. 1929.

Cyathea obtusa (Kaulf.) Domin, Pteridophyta 264, 1929.

Cyathea obtusa var. bullata (Christ) Domin, Acta Bot. Bohem. 9: 142. 1930.

48 FIELDIANA: BOTANY, VOLUME 37

Leaves terminating abruptly in a pinnatifid apical section; petioles with stout

spines to 3 mm. long; rachis and costae beneath with brown (or very rarely a

few whitish) scales; pinnae cut three-fourths to seven-eighths to the costa;

costules with glossy, amber to castaneous, bullate scales occasional to abundant

beneath, or very rarely some whitish or bicolorous scales intermingled; secondary

segments falcate, obtuse; veins once- (rarely twice-) forked; sori in a single line

between costule and segment margin.

No type was designated by Kaulfuss for Hemitelia obtusa. However,

Krug (1898) cites a Ryan specimen from Montserrat as well as Sieber

ft. mixta No. 331 (no locality) as H. obtusa from Kaulfuss' herbarium.

It is likely these are the specimens that provided the basis for Kaulfuss'

description. Hence these specimens (destroyed at LZ) can be con-

sidered as syntypes. I have seen no duplicates of them.

Of hundreds of specimens of Cnemidaria grandijolia examined, one

juvenile plant collected in 1938 on Morne Gimie, St. Lucia (Box 7963)

fails to fit the pattern of distribution. Though the leaves are too imma-

ture for positive determination, the brown and rather glossy scales of

the axes lead to a tentative identification as var. obtusa. Furthermore,

a few specimens of var. obtusa from St. Vincent have been observed with

a scattering of very light colored — nearly white — scales interspersed

among the brown, so they appear similar to those of var. grandifolia.

Thus it seems that St. Vincent and St. Lucia, where the ranges of the

varieties converge, comprise an "area" where they also tend to inter-

grade.

In and at edges of forests, on stream banks, and mountainsides, 300-

1,050m. St. Vincent, Grenada, Tobago, Trinidad, Venezuela (Margarita

Is., Paraguana Peninsula) .

Selected specimens examined. — St. Vincent: In glens, about 1,000

ft., H. H. & G. W. Smith 854 (GH, IJ, MO, US). Rain forest, Beard

1434 (F, MO). Morne Garou Mountains, Morton 5072 (GH, MO, NY,

US). Grenada: Forest borders, Grand Etang, Beard 1194 (F, GH, US).

Above Windsor Forest, St. David Parish, Proctor 16904 (GH, IJ, U,

US). Tobago: Eggers 5859 (F, US). Trinidad: Parker s.n. (GH). Ven-

ezuela: San Juan Mountain, Margarita Island, Johnston 191 (F, GH,

NY, US). — some specimens a mixture with C. spectabilis var. specta-

bilis. Cerro Santa Ana, Paraguana Peninsula, Curran & Haman 676 &

736 (GH) and 683 (GH, US).

8. Cnemidaria bella (Mett.) Tryon. Contr. Gray Herb. 200: 52.

1970. Figure 16.

Hemitelia bella Mett., Fil. Hort. Lips. 110. 1856. TYPE COLLECTION: "Hort.

Bot. Lips., cult." AUTHENTIC SPECIMENS: K! P! MO! US!

STOLZE: REVISION OF CNEMIDARIA

49

FIG. 16. Cnemidaria bella, T. C. Porter s.n. (U.S. Herb. No. 2204421), cult.

Bot. Garden, Leipzig.

Cyathea bella (Mett.) Domin, Pteridophyta 263, 1929.

Caudex, petiole and leaf apex not seen; leaves to 1 m. broad; rachis non-alale,

lacking scales and trichomes; pinnae distant, stalked (to 2 cm.), cut one-third

to a little more than halfway to the costa; costae and costules with scales and

trichomes lacking, except for rare, dull brown, amorphous scales on the costae

beneath; secondary segments subfalcate, broadly rounded; veins simple to once-

forked, commonly free but occasionally some basal ones merging to form costal

areoles, basal basiscopic ones usually arising from the costule or, rarely, from the

costa; sori in a single line, submarginal on the segments; indusia dark brown,

semicircular, with 2 to 3 lobes.

Hooker's (1865) description mentions "st. aculeated at the base;"

i.e., the petiole spiny at the base. Although no petioles were available

in material I examined, portions of the rachis were strongly muricate, a

condition generally occurring when petioles are spiny.

Cnemidaria bella is a highly distinctive species, not likely to be con-

fused with any other, but it is somewhat questionable that it be included

50 FIELDIANA: BOTANY, VOLUME 37

as a valid species. The original collections were from cultivated ma-

terial at the Botanical Garden at Leipzig, and apparently the species

has never been found in the wild. The sum total of its representation

today are the fragments and parts of leaves which have made their way

into various herbaria. The few mature spores seen have the three, large,

characteristic Cnemidarioid pores, the lamina is completely devoid of

trichomes, and the veins, though mostly free, are connivent to the sinus,

so the species must definitely be placed in Cnemidaria. However, the

pinnae are relatively long-stalked, a condition otherwise unknown in the

genus, the soral line is submarginal (rare in the genus), and most spores

are abortive. All of these characters indicate that the plant at the Leipzig

Gardens may have been a hybrid between a species of Cnemidaria and

one of Cyathea. One likely parent would be Cyathea speciosa or C.

integrifolia, both of which have subentire or entire, short-stalked pinnae,

submarginal sori, and trichomes thinly scattered on costae and rachis.

The Cnemidaria parent might be C. horrida or C. spectabilis, both com-

mon species, with supramedial to submarginal sori, costal areoles, and

indument rare to lacking.

9. Cnemidaria uleana (Samp.) Tryon, Contr. Gray Herb. 200: 52.

1970.

Caudex to 0.5 m. long and 3 cm. in diameter; leaves to 2 m. long and 0.7 m.

broad, pinnatifid apical section gradually reduced (or conform in var. abita-

guensis); petiole to 0.7 m. long, smooth to slightly muricate, the scales bicolorous,

dark brown with broad whitish margins, linear-lanceolate to lance-ovate, trichomes

lacking; rachis non-alate, scales sparse, pale to bicolorous or lacking, trichomes

lacking; pinnae sessile, cut two-thirds to seven-eighths to the costa, basal pair dis-

tinctly reduced and often deflexed; costae and costules with broad, amorphous,

pale yellowish or whitish scales scattered to rare beneath, trichomes lacking;

secondary segments falcate to subfalcate, obtuse to acute, basal pair often over-

lapping the rachis; veins once-to thrice-forked, free, or basal ones often merging

to form costal areoles, basal basiscopic ones (especially in basal one-third or

apical one-third of pinna) arising from the costa; sori in one or two lines, medial

to supramedial between costule and segment margin; indusia pale yellow or

yellowish brown, semicircular, subentire to several-lobed.

Cnemidaria uleana appears to have been derived from the same stock

as C. horrida, which shares the range of var. uleana in Peru and that of

var. abitaguensis in Colombia and Ecuador. The typical variety of

C. uleana is the sole representative of the genus in Brazil. C. uleana var.

uleana exhibits a condition unique in the genus, in that the degree of

anastomosing of basal veins is inconstant, i.e., one leaf may be pre-

dominantly free-veined, another mostly areolate along the costa, and

STOLZE: REVISION OF CNEMIDARIA 51

yet another may show an equal mixture of both conditions. Var.

abitaguensis is always free-veined.

9a. Cnemidaria uleana var. uleana. Figure 9b, Map 8.

Hemitelia uleana Samp., Bol. Mus. Nac. Rio de Janeiro 1: 65. 1923. TYPE

COLLECTION: Vie s.n., Perto de Nova Friburgo, Alto da Serra. Est. do Rio,

Brazil, 1898. HOLOTYPE: R! PHOTO: F! GH!

Hemitelia maxonii Rosenst., Repert. Spec. Nov. Regni Veg. 21: 344. 1925.

TYPE COLLECTION: Brade 6901, Serra do Mar. Est. Sao Paulo, Brazil, 26/1 V/

1914. HOLOTYPE: R! ISOTYPE: S-PA!

Cyathea subarborescens Domin, Acta Bot. Bohem. 9: 162. 1930, nom.nov.

for Hemitelia maxonii Rosenst., not Cyathea maxonii Underw. ex Maxon.

Leaves terminating in a gradually reduced, pinnatifid apical section; rachis

with pale to bicolorous scales scattered beneath; scales of costae and costules

beneath scattered, whitish, sometimes faintly red-margined.

Specimens of C. uleana var. uleana with costal areoles could be con-

fused with C. speciosa in Peru, but the latter has pinnae lobed less than

halfway to the costa and leaves terminating in conform or subconform

apical sections. C. tryoniana and some varieties of C. mutica are some-

what similar to free-veined specimens of var. uleana, but are distin-

guished by the characters shown in the key.

Until recently plants of the typical variety of C. uleana were known

only from Brazil, but some fine collections were made by Dudley in

1968 from Dept. Cuzco, Peru. However, it is unlikely that further

collections will be made between these two disjunct regions (except

perhaps in the wet, montane areas of Bolivia contiguous with Peru)

since the combinations of altitude and climate favorable to the growing

habits of Cnemidaria are lacking.

Forests, shaded ravines, and hillsides, 580-2, 000m., Brazil, Peru.

Selected specimens examined. — Peru. Cuzco: Prov. La Convention,

Dudley 10459, 11312 (GH, NA). Brazil. Minas Gerais: 2-3 km. ENE

of Ouro Preto, Tryon & Tryon 6871 (GH). Parana: Serra do Mar,

Dusen 17306 (F, GH, MO, NY). Rio: Rio Bonito, Itatiaia, Pereira 312

(F, GH, MO, US). Sao Paulo: Serra do Bocania, Lutz 727 (US). Alto

da Serra, L. B. Smith 2077 (GH, NY, R, US).

9b. Cnemidaria uleana var. abitaguensis (Domin) Stolze, stat. et

comb. nov. Map 8.

Hemitelia abitaguensis Domin. Mem. Roy. Czech. Soc. Sci. 2: 74. 1929. TYPE

COLLECTION: Spruce 5364, Monte Abitagua, Andibus Ecuadorensibus. HOLO-

TYPE: K! PHOTO: F! ISOTYPES: GH! NY!

Cyathea abitaguensis (Domin) Domin, Acta Bot. Bohem. 9: 88. 1930.

52 FIELDIANA: BOTANY, VOLUME 37

Leaves terminating in a conform or subconform apical pinna; rachis lacking

scales; Costae and costules with pale to light brown scales very rare beneath, or

lacking.

Other than the character used in the key, little can be seen to separate

C. uleana var. abitaguensis from the free-veined specimens of the typical

variety, except that scales are virtually absent on the axes of the latter,

and the rare scales found are pale yellowish or very light brown. Scales

of var. uleana, though not abundant, can be found scattered along the

axes, and are generally whitish, with those of the rachis sometimes pos-

sessing a thin, dark brown median stripe, or those of the costae and cos-

tules occasionally with a thin, reddish margin. Such subtle differences

would not merit even varietal distinction, but they are reinforced by

the character of the apical section. Future collections of var. abitaguen-

sis will confirm the value of this character or demonstrate intergradation

with var. uleana.

Forests, 1,900-2,100 m., Colombia and Ecuador

Additional specimen examined. — Colombia. Huila: Hondonada del

Abra de San Andres, Cuatrecasas 8640 (F, US) .

10. Cnemidaria cocleana Stolze sp. nov. Figure 17, Map 12.

Petiolus spinis usque ad 3 mm. longis instructus; pinnae per mediam plus

minusve distantiam inter apices pinnularum et costam incisae; pinnae basales

aliquantum deminutae et deflexae; venae 1- vel 4-furcatae, venae infimae anas-

tomosantes et areolas costales efformantes; venae infimae basiscopicae e costulis

egredientes; sori uniseriati, inter costulam et marginem medii vel inframediani;

indusia omnino circularia.

TYPE COLLECTION: La Mesa, 5 miles N. El Valle, Code, Panama, Nov.

10, 1965, Tyson, Godfrey et al. 2452. HOLOTYPE: (3 sheets) FSU! PHOTO: F!

GH!

Caudex not seen; leaves to 0.8 m. broad, terminating in an abruptly reduced

apical section; petiole with spines to 3 mm. long, the scales sparse, bicolorous,

dark brown with whitish margins, trichomes lacking; rachis non-alate, the scales

rare, bicolorous, the minute, recurved trichomes sparse to scattered beneath;

pinnae sessile, incised more or less halfway to the costa, basal pair somewhat

reduced and deflexed; costae and costules without scales or trichomes; secondary

segments falcate, obtuse, basal pair commonly overlapping the rachis; veins 1-

to 4-times forked, basal ones merging to form costal areoles, basal basiscopic ones

arising from the costules; sori in a single line, medial to scarcely inframedial be-

tween costule and segment margin; indusia yellowish brown, fully circular, with

margins subentire and undulate.

This species bears a resemblance to C. spectabilis and C. choricarpa;

however, both have the pinnae lobed consistently less than halfway to

the costa, and their indusia have the conventional semicircular shape,

whereas Cnemidaria cocleana has at least its larger pinnae cut more

FIG. 17. Cnemidaria cocleana, Tyson et al. 2452 (FSU, holotype).

53

54

FIELDIANA: BOTANY, VOLUME 37

than halfway to the costa and the indusia forming an unbroken circle

(fig. lie) around the receptacle. Furthermore, C. cocleana does not have

the broadly alate rachis of C. ctwricarpa. The completely circular in-

dusium is a rather unique character in the genus, otherwise being found

only in C. nervosa, the species from Peru and Ecuador with subentire

pinnae.

The species has been collected in the deep shade of forests, only in

the province of Code, Panama, 760-1 ,000 m.

Additional specimen examined. — Panama. Code: Crater of El Valle

de Anton, La Mesa, Wilbur etal. 11,111. (F, NY).

11. Cnemidaria quitensis (Domin) Tryon, Contr. Gray Herb. 200:

52. 1970. Figure 18, Map 11.

Hemitelia quitensis Domin, Kew Bull. 215, 1929 (non Cyathea quitensis

[C.Chr.] Domin, 1929). TYPE COLLECTION: Sodiro s.n., In Ecuador in Andi-

bus Quitensis, 1875. HOLOTYPE: K?

Cyathea andicola Domin, Acta Bot. Bohem. 9: 91, 1930; nom. nov. for Hemi-

telia quitensis Domin.

Caudex to 2 m. long and 4 cm. in diameter; leaves to 3 m. long and 1 m. broad,

terminating in a conform apical pinna; petiole to 1.2 m. long, with spines to 3 mm.

long, the scales sparse, bicolorous, dark brown with whitish margins, mostly

ovate; rachis non-alate, scales sparse, pale to bicolorous or lacking, the minute,

terete, recurved trichomes abundant beneath; pinnae subsessile to sessile, cut one-

third to a little more than halfway to the costa, the basal pair somewhat reduced;

costae and costules with broad, amorphous, pale to tawny scales scattered be-

neath, the trichomes scattered to lacking beneath; secondary segments falcate

to subfalcate, obtuse, commonly imbricate with their sinuses cartilaginous above

and contrasting in color with adjacent tissue, basal segments often overlapping the

rachis; veins once- to thrice-forked, basal ones commonly merging to form costal

areoles, basal basiscopic ones commonly arising from the base of the costule, or

FIG. 18. Cnemidaria quitensis, Ewan 16812 (US): portion of pinna, adaxial

side, showing imbricate segments and cartilaginous sinuses.

STOLZE: REVISION OF CNEMIDARIA 55

infrequently from the costa; sort in one or two lines, commonly inframedial be-

tween costule and segment margin; indusia pale yellow or yellowish-brown, more

or less semicircular, subentire to several-lobed.

When a loan for this study was prepared at Kew, the holotype of

H. quitensis was not located, although it would be expected to be there

along with other Domin types.

Pinnae of Cnemidaria quitensis are commonly cut halfway or less to

the costa, but in some of the larger specimens a number of pinnae may

be incised more deeply. The less deeply divided specimens might be

confused with C. spectabilis var. colombiensis, especially when the leaf

apex is not available. However, the soral line in C. quitensis is consis-

tently nearer to the costule than the margin, the minute, terete, recurved

trichomes are scattered to abundant on the rachis beneath, and there

are frequent whitish or pale to tawny scales on the costae and costules

beneath. In C. spectabilis var. colombiensis, the soral line is usually

situated midway between costule and margin, trichomes are lacking, and

the rare scales on the axes beneath are commonly dull and dark brown.

The cartilaginous sinuses of C. quitensis afford a good diagnostic char-

acter within the genus (fig. 18), since only C. spectabilis var. colom-

biensis and, to a lesser degree, C. cruciata exhibit this condition. Es-

pecially evident on the adaxial side, the veins converge so strongly at

the sinuses as to form a cartilaginous mass, yellowish or very light

brown in color in dried material, which contrasts markedly with the

adjacent tissue.

In forests, Colombia and Ecuador, 100-1 ,400 m.

Selected specimens examined. — Colombia. Caldas. Cordillera Occi-

dental, Santa Cecilia, von Sneidern 5199, (F, US). Cauca: La Gallera,

Micay Valley, Killip 7774, (GH, NY, US). Choco: N. of San Jose del

Palmar, Lellinger and de la Sota 728, (US). Narino: Near Altiquer, Rio

Guabo, Ewan 16812 (GH, US); Barbacoas, Corregimento Santander a

Barbacoas, Garcia-Barriga 13120 (US); Forest, Ricaurte, von Sneidern

A. 473 (GH). Ecuador. Esmeraldas: Playa Rica, Parroquia de Con-

ception, Mexia 8479 (F, GH, US) ; El Oro: Near Moromoro, 21Vi miles

west of Portovelo, Wiggins 10930 (MO, US). Imbabura: Entre El Pajon

y Cachaco, Acosta Soils 12720 (F).

12. Cnemidaria alatissima Stolze sp. nov. Figure 19, Map 11.

Petiolus spinis brevibus et paleis numerosis latis albidis instructus; rachis ex

toto alata; pinnae sessiles, per duas tertias vel tres quartas partes distantiae inter

apices pinnularum et costam incisae; venae furcatae vel bifurcatae, venae infimae

anastomosantes et areolas costales efformantes; venae infimae basiscopicae e

costulis egredientes; sori uniseriati, inter costulam et marginem medii.

PLANTS OF

FIG. 19. Cnemidaria alatissima, Dudley 13282 (GH, holotype).

56

STOLZE: REVISION OF CNEMIDARIA 57

TYPE COLLECTION: Dudley 13282, Rio Llulla Pichis watershed, Dept.

Huanuco, Peru, 25 July 1969. HOLOTYPE: (2 sheets) GH! PHOTO: F!

ISOTYPE: NA!

Caudex short or lacking; leaves to 2 m. long and 0.5 m. broad, terminating in

a subconform apical pinna; petiole to 1.5 m. long with minute spines, thickly

covered near the base with large, ovate, whitish scales; rachis provided on either

side with a green wing 1 to 2 mm. wide, and broad amorphous, whitish scales

scattered beneath, trichomes rare to scattered beneath; pinnae sessile, cut two-

thirds to three-fourths to the costa, basal pair not seen; costae and costules lacking

scales or trichomes; secondary segments falcate, obtuse to acute, the basal pair

often overlapping the rachis; veins once- to twice-forked, basal ones commonly

merging to form costal areoles, basal basiscopic ones usually arising from the

costule; son in a single line approximately midway between costule and segment

margin; indnsia brownish, often rudimentary, usually less than semi-circular, sub-

entire.

The broadly alate rachis and the large white scales of the petiole

make this species one of the most striking in the genus. Green wings to

2 mm. wide extend along either side of the rachis from apex to base,

and even well down the petiole. The lower part of the petiole is nearly

obscured by the abundant scales, which measure up to 1.5 cm. long

and 0.7 cm. broad. These are commonly creamy-white throughout, but

some at the very base of the petiole have a thin, shiny-brown median

stripe at their tips. The indusia, also, are rather distinctive. Although

characteristically hemitelioid in that they are nearly semicircular in out-

line, they are so small and thin and so appressed to the leaf tissue that

they can be observed only by scraping away most of the sporangia. The

only other species of Cnemidaria possibly confused with this would be

C. choricarpa of Costa Rica, especially in that both have strongly alate

rachises. But C. alatissima can be easily distinguished by its much deeper

lobing, as well as by the abundant whitish scales.

Thus far known only from the type collection, and one juvenile plant

(Dudley 13281) collected with the type; found in large numbers in dark

cloud forest, in wet places, especially in spray of waterfall, Huanuco,

Peru, ca. 1,540 m.

13. Cnemidaria choricarpa (Maxon) Tryon, Contr. Gray Herb. 200:

51,1970. Figure 2, Map 12.

Hemitelia choricarpa Maxon, Contr. U.S. Natl. Herb. 16: 40. 1912. TYPE

COLLECTION: Pinter 4835, Buenos Aires, Costa Rica. HOLOTYPE: US!

PHOTO: F! GH!

Cyathea choricarpa (Maxon) Domin, Pteridophyta 263, 1929.

Caudex to 0.6 m. long and 2.5 cm. in diameter; leaves to 2 m. long and 0.6 m.

broad, terminating in an abruptly reduced, pinnatifid apex; petiole to 0.8 m. long,

58 FIELDIANA: BOTANY, VOLUME 37

spines to 3 mm. long or, rarely, lacking, the lanceolate scales bicolorous, dark

brown with narrow, whitish margins, sparse except at the very base; rachis pro-

vided on either side with a narrow green wing, up to 1.5 mm. wide above, often

vestigial below, the scales dark brown, scattered or lacking, the minute, terete,

recurved trichomes scattered to mostly abundant beneath; pinnae sessile to sub-

sessile, cut one-third to halfway to the costa, basal ones somewhat reduced and

occasionally deflexed; costae and costules with broad, brown, amorphous scales

scattered beneath, and trichomes scattered to lacking beneath; secondary seg-

ments falcate, obtuse, basal pair crowding or overlapping the rachis; veins simple

to once-forked, basal ones mostly merging to form costal areoles, basal basiscopic

ones arising from the costule; son in a single line, medial to mostly inframedial

between costule and segment margin; indusia brownish, more or less semicircular,

subentire to erose, or slightly lobed.

Cnemidaria choricarpa is closely related to C. mutica of Costa Rica

and Panama and C. decurrens of Mexico and Guatemala, sharing a num-

ber of morphological features with both, and intermediate between

them in other characters. The winged rachis is common in C. decurrens

and is frequently found in C. mutica, at least in vars. grandis and

chiricana. Furthermore, all three species are similar in the shape of

their leaf apices and vestiture. The subentire or shallowly lobed pinnae

of C. decurrens are commonly areolate along the costae, but those of

C. mutica are quite deeply lobed and free-veined. Thus C. choricarpa lies

midway between these species in having pinnae cut one-third to halfway

to costa, and although the basal veins generally merge to form costal

areoles, many specimens can be found which show occasional tendencies

to free venation. In fact, Scamman 5881 from San Isidro and L. O.

Williams et al. 28785 from Rio Sonador are notable in their almost

complete lack of costal areoles.

The range of C. choricarpa is basically confined to Costa Rica, but two

collections that probably represent the species have been seen from

Colombia. A recent collection from Choco. Lellrnser & de la Sota 277,

matches C. choricarpa in every way, except that it lacks petiole spines.

If subsequent collections lacking spines are found from this area, then

perhaps it will merit varietal distinction. One other collection from

Colombia, Triana s.n., is mounted on a sheet at Kew, together with a

pinna of C. quitensis, and is apparently C. choricarpa, but unfortunately

no petiole is available to examine. Hence, on the strength of these two

collections, the range of the species is provisionally extended to Colom-

bia.

At least two sheets of Pittier 10969 are extant. I have examined one —

the holotype of Hemitelia pittieri (US); the other is Cnemidaria chori-

carpa (CR). A discussion of this mixed collection may be found under

C. mutica var. mutica.

STOLZE: REVISION OF CNEMIDARIA 59

Wet, forest slopes, Costa Rica and Colombia, 30-1 ,700 m.

Selected specimens examined. — Costa Rica. Puntarenas: Slopes above

Golfito, Burger & Matta 4756 (F). 4 miles west of Rincon, Osa Penin-

sula, Burger & Stolze 5499 (F, GH). Canas Gordas, Pittier 10966, (US)

and 70969 p.p. (CR). San Jose: Vicinity of San Isidro El General,

Molina et al 17954 (CR. F, GH. MO). Vicinity of El General, Skutch

2265 (GH, MO, US). Colombia. Choco: N.W. side of Alto del Buey,

Lellinger & de la Sot a 277 (COL, CR, F, HUA, LP, US). Narino: Entre

Tuquerres et Barbacoas, Triana s.n. (K).

14. Cnemidaria chocoensis Stolze sp. nov. Figure 20, Map 12.

Petiolus baud spinosus; pinnae sessiles, per unam tertiam vel mediam partem

distantiae inter apices pinnularum et costam incisae; pinna apicalis subconformis,

segmentis basalibus elongatis; costae et costulae subtus paleis atrobrunneis,

nitidis instructae; venae 2- vel 3-furcatae, venae infimae anastomosantes et

areolas costales efformantes; venae infimae basiscopicae e costis egredientes; sori

uniseriati, inter costulam et marginem manifeste inframediani.

TYPE COLLECTION: Lellinger & de la Sola 763, 2 km. east of San Jose

del Palmar, Choco, Colombia. HOLOTYPE: US! PHOTOS: F! GH! ISOTYPES:

COL, CR, F! HUA, LP.

Caitdex not seen; leaves to 2.3 m. long and 0.4 m. broad, terminating in a sub-

conform apical pinna, this usually with a pair of elongate basal lobes; petiole

to 1.2 m. long, spines lacking, the scales scattered to frequent, bicolorous, dark

brown, shiny, with broad pale to whitish margins, trichomes lacking; rachis non-

alate, the scales scattered to frequent, dark shiny-brown or rarely with whitish

margins, trichomes lacking; pinnae sessile, cut one-third to halfway to the costa,

basal ones slightly reduced; costae and costules with broad, dark shiny-brown

scales scattered to abundant beneath, trichomes lacking; secondary segments sub-

falcate, obtuse, basal pair crowding or overlapping the rachis; veins 2- to 3-times

forked, basal ones commonly merging to form costal areoles, basal basiscopic

ones (especially in the basal one-third or apical one-third of pinna) arising from

the costae; sori in a single line, distinctly inframedial between costule and seg-

ment margin; indusia pale to yellowish brown, more or less semicircular, sub-

entire to shallowly lobed.

This species has the lamina terminated by a subconform, apical

pinna, like many specimens of C. speciosa, i.e., the terminal section

closely resembles the larger pinnae in size and shape except for a

pair of rather elongate basal segments. C. chocoensis is similar to C.

speciosa in many other respects, but may be distinguished from the

latter by its obviously inframedial sori, the frequent, shiny brown scales

along the axes, and the basal basiscopic veins arising mainly from the

costa. C. speciosa has medial to supramedial sori, the scales along the

axes, when present, are dull brown, and the basal basiscopic veins arise

•

FIG. 20. Cnemidaria chocoensis, Lellinger & de la Sota 763, (US, holotype).

60

STOLZE: REVISION OF CNEMIDARIA 61

from the costules. C. chocoensis could also be confused with C. ewanii,

but may be easily separated by the characters noted in the key.

Another Choco specimen seen, Lellinger & de la Sola 853, is identical

with the type in every way except that its pinnae are much thicker in

texture and appear rather constricted, i.e., much narrower in relation

to their length, and they are a dark brown color when dry instead of

olive green. Perhaps it is merely an altitudinal variant, but is at least

provisionally determined as C. chocoensis along with the other two

collections from the area.

Disturbed secondary forests, Choco, Colombia, 750-1, 650m.

Additional specimens examined. — Colombia. Choco: Along Bolivar-

Quibdo road, Lellinger & de la Sota 895 (COL, CR, F, HUA, LP, US).

8 km. S.W. of El Cairo, Lellinger & de la Sota 853 (COL, CR, HUA,

LP,US).

15. Cnemidaria spectabilis (Kunze) Tryon, Contr. Gray Herb. 200:

52. 1970.

Caudex to 0.5 m. long and 5 cm. in diameter; leaves to 1.7 m. long and 0.8 m.

broad, terminating in an abruptly reduced, pinnatifid apex; petiole to 1.2 m. long,

spines to 5 mm. long, the scales sparse or lacking except at the very base, lanceo-

late to linear-lanceolate, bicolorous, dark brown with narrow whitish, often fim-

briate margins; rachis often spiny toward base of lamina, non-alate, except rarely

between the several upper pinnae, scales and trichomes usually lacking; pinnae

sessile, cut one-third to halfway to the costa, basal ones somewhat reduced and

deflexed; costae and costules with brown, broad, amorphous scales scattered or

lacking beneath, trichomes lacking; secondary segments subfalcate, obtuse to rare-

ly subapiculate, basal pair crowding or overlapping the rachis; veins simple to

thrice-forked, basal ones commonly merging to form costal areoles, basal basi-

scopic ones arising from the costule; sori in a single line, medial to supramedial

between costule and segment margin; indusia pale to yellowish-brown, more or

less semicircular (rarely almost fully circular in var. colombiensis) , subentire to

lobed.

Lying approximately midway along the phyletic line, this species ranks

next to C. horrida as the most widespread species in the genus. Speci-

mens have been collected all along the northern edges of South America,

from Colombia through the Guianas, including Trinidad and Tobago. It

is likely that seven other species, from northern South America to

Bolivia, have been derived from C. spectabilis or its ancestral stock (see

fig. 1 ) , and all may be considered "typically Cnemidarioid," in that

their pinnae are strongly areolate and rather shallowly lobed.

15a. Cnemidaria spectabilis var. spectabilis Figures 3b, lOb; Map 9.

Hemitelia spectabilis Kunze, Linnaea 21: 233. 1848. TYPE COLLECTION:

62 FIELDIANA: BOTANY, VOLUME 37

Kappler 1771, Surinam, pr. stationem Victoriam, Apr. 1847. ISOTYPES: K!

P! PHOTO: F!

Actinophlebia obtusa Presl, Gefassb. Stipes der Farm 48, 1847 (preprint of

Abh. Bohm Ges. Wiss. V. 5: 356. 1848) (non Hemitelia obtusa Kaulf. 1824.).

Based on Hooker's plate 14, Sp.Fil. 1. 1844.

Hemistegia spectabilis (Kunze) Fee, Mem. Fam. Foug. 5 (Gen.Fil.): 351.

1850-52.

Hemitelia klotzschiana Klotzsch, Allg. Gartenzeitung 20: 42. 1852. TYPE

COLLECTION: probably Karsten 143, Venezuela, Jan. 1852. PROBABLE ISO-

TYPE: B! PHOTO: F!

Hemitelia spectabilis var. trinitensis Domin, Mem. Roy. Czech. Soc. Sci. 2:

72. 1929. TYPE COLLECTION: Wake field s.n., La Seiva Valley, Trinidad,

16/1/1921.

Hemitelia spectabilis var. longipinna Domin, tom.cit. 73. (Domin's "typical

variety", based on H. spectabilis Kunze.)

Cyathea spectabilis var. trinitensis (Domin) Domin, Pteridophyta 264, 1929.

Cyathea spectabilis var. longipinna (Domin) Domin, loc. cit.

Larger pinnae 3-5 (6) cm. wide and 28-36 (38) cm. long, secondary segments

approximate or divergent, only rarely subimbricate, the sinuses rarely (if ever)

cartilaginous, the tissue more or less uniformly green; soral line mostly supra-

medial between costule and segment margin; indusia more or less semicircular.

The typification of Hemitelia klotzschiana Kl. is somewhat in doubt.

Although no type is designated in the original description, Klotzsch's

citation of "Karsten" after the name, implies the type is from a Karsten

collection. There is a specimen of Karsten 143 at Berlin marked "Hem-

itelia (Cnemidaria) klotzschiana Karsten" which could very well be

the plant Klotzsch had in mind.

This variety of the species most closely resembles Cnemidaria con-

similis and C. cruciata. From the latter, it may be distinguished by the

characters of leaf apex and pinna segments noted in the key. It also dif-

fers from C. consimilis in shape of leaf apex, but, unfortunately, this

feature is often lacking on many herbarium sheets, and in these in-

stances the position and color of the petiole scales furnish the most re-

liable diagnostic character. In C. spectabilis var. spectabilis the scales

are rare or lacking on the petiole, except at the very base. They are

bicolorous, i.e., mostly dark brown with very narrow whitish margins;

however, in C. consimilis the scales are either completely white or with

a very narrow brown median stripe, and are quite abundant throughout

the petiole, from lamina to base.

Two Steyermark specimens, 89525 and 92924 from Estado Bolivar,

Venezuela, are especially close to C. consimilis, because of the some-

what subconform apical pinna. However, the petiole scales are dark

brown, with very narrow whitish margins, and are present only at the

base, as in typical C. spectabilis.

STOLZE: REVISION OF CNEMIDARIA 63

In and at the edges of forests, along stream banks and mountain-

sides, 100-1,200 m., Venezuela, the Guianas, Trinidad, and Tobago.

Selected specimens examined. — Trinidad: Balandra Bay, E. G. Brit-

ton et al. 433 (GH, NY, US). Shaded gully, Dibe Valley, N. L. Britton

et al. 1748 (F, GH, NY, US). 1877-78 (no locality), Fender 25 (F,

GH, MO, NY, US). Tobago: Main ridge, Broadway 3886 (F, GH, MO,

NY, U. US). Near Caledonia, Broadway 4540 (F, GH, MO, NY, U,

US). Venezuela: Lower Orinoco, Eleanor Creek, Rusby & Squires 110

(F, GH, MO, NY, US). Anzoategui: N.E. of Bergantin, by Sucre

boundary, Steyermark 61207 (F, US, VEN). Merida: prope Coloniam

Tovar, Fendler 480 (GH, K, MO). Miranda: Cardenas, Siquire Valley,

Pittier 7091, (GH, US, VEN) and 5945 (F, NY, US). Nueva Esparta:

Margarita Island, El Valle, Miller & Johnston 164 (F, GH, MO, US).

Sucre: Aricagua, vicinity of Cristobal Colon, Broadway 567 (GH, NY,

US). British Guiana: Demerara, Region Mt., 1895, Jenman s.n. (NY).

Demerara, Isorooroo River, 1899, Jenman s.n. (NY). Surinam: Rika-

nau, near Moengo, Lindeman 6124 (K, MO, U. US). Distr. Broko-

pondo, trail from Bronsweg, Kramer & Hekking 3195 (U). French

Guiana: "Guyane," coll. unknown, F herbarium No. 808351 (F).

Guyana, in sylvis humidis, LePrieur s.n. (K).

15b. Cnemidaria spectabilis var. colombiensis Stolze, var. nov. Figure

21, Map 9.

Pinnae maximae 5-7 cm. latae, 34-42 cm. longae; segmenta secundaria plerum-

que imbricata vel subimbricata, sinibus plerumque cartilagineis et colore suo con-

trastantibus cum textura adiacente; sori inter costulam et marginem medii; indusia

semicircularia vel fere omnino circularia.

Largest pinnae 5-7 cm. wide and 34-42 cm. long; secondary segments commonly

imbricate or subimbricate (at least near apices), the sinuses mostly cartilaginous

above and contrasting in color with adjacent tissue; soral line medial between

costule and segment margin; indusia semicircular to almost fully circular.

TYPE COLLECTION: Killip 5039, Cordoba, Dept. El Valle, Colombia, May

6-8, 1922. HOLOTYPE: (3 sheets) US! PHOTO: F! ISOTYPES: GH! NY!

Besides the characters noted in the key, this variety can often be sep-

arated from var. spectabilis by its wider pinnae. Although some rare

specimens of the latter can be found which have occasional pinnae to

6 cm. wide, the largest pinnae normally measure from 3-5 cm. wide,

whereas the largest pinnae of var. colombiensis commonly measure 6-7

cm. wide.

The indusia of var. colombiensis are commonly much more than

semicircular in shape, occasionally appearing to be almost fully circular,

FIG. 21. Cnemidaria spectabilis var. colombiensis, Killip 5039, (US, holotype).

64

STOLZE: REVISION OF CNEMIDARIA 65

such as in C. cocleana and C. nervosa. But closer examination will re-

veal a sinus, which is sometimes partially obscured by the overlapping

tissue of the indusium.

Wooded areas, Colombia, 70-500 m.

Additional specimens examined. — Colombia. Choco: Between La

Oveja and Quibdo, Archer 1764 (US). Second growth forest, Anda-

goya, Killip 35377 (GH, US). West of Istmina, road to Pie de Pepe,

Lellinger & de la Sota 431 (US).

16. Cnemidaria cruciata (Desv.) Stolze. comb. nov. Figure 3a, Map

9.

Hemitelia cruciata Desv., Prodr. 320. 1827. TYPE COLLECTION: Desvaux

s.n., "Habitat in America." HOLOTYPE: P! PHOTO: F! GH!

Hemitelia leprieurii Kunze, Bot. Zeit. II: 296. 1844. TYPE COLLECTION:

LePrieur 200, Fr. Guiana.

Cyathea leprieurii (Kunze) Domin, Pteridophyta 264. 1929.

Caudex not seen; leaves to 0.6 m. broad, reduced very gradually to a pinnati-

fid apex, lacking a distinct apical section; complete petiole not seen, spines to

2 mm. long, scales lacking on most of petiole seen; rachis often short-spiny

toward the base, non-alate, scales and trichomes lacking; pinnae sessile, or lower

ones sometimes short-stalked, cut one-third to halfway to the costa, basal ones

somewhat reduced and deflexed; costae and costules without scales or trichomes;

secondary segments subfalcate, obtuse to occasionally subapiculate, commonly

imbricate, with their sinuses tending to be cartilaginous above, basal segments

often crowding or overlapping the rachis; veins simple to twice-forked, basal ones

commonly merging to form costal areoles, basal basiscopic ones arising from the

costule; sort in a single line, supramedial between costule and segment margin;

indusia pale to yellowish-brown, more or less semicircular, subentire to lobed.

This species is poorly represented in herbaria, and I have not seen

a caudex, or even a complete petiole. The type consists only of a rachis

with four pairs of pinnae, and most other material is even more frag-

mentary. Only one specimen shows the leaf apex.

It has been suggested that Cnemidaria cruciata is equivalent to C.

spectabilis. Indeed, they are very closely related; but the aspect of the

pinnae, segments, and leaf apices is quite distinct in each species, and

if more complete collections are made in the future, perhaps additional

differences will be noted. The pinnae of C. spectabilis are rather widely

separated, while those of C. cruciata are approximate and, at least near

the leaf apex, often crowded or even overlapping. The leaf apex of C.

cruciata is like no other in the genus. An excellent specimen (LePrieur,

1838) on four sheets at the U.S. National Herbarium reveals a leaf

gradually reduced to the tip, so that it lacks a distinct terminal section.

This is in marked contrast to all other species of Cnemidaria, which have

66 FIELDIANA: BOTANY, VOLUME 37

their leaves abruptly reduced into a conspicuous, separate apical section,

or terminating in a conform terminal pinna (see fig. 3).

According to the labels, all specimens of Cnemidaria cruciata ex-

amined are from French Guiana. Only Desvaux's type is in doubt, for

the label is marked simply "Habitat in America." But although the

source of his specimen may not be known, there is no evidence to sug-

gest that it was collected anywhere other than French Guiana.

Additional specimens examined. — French Guiana: Guyane Francaise,

1838, LePrieur s.n. (US). Guyane, Jul. 1824, Poiteau s.n. (K). Guiana

Francaise, 1824, Poiteau 139 (B, US). Acarouany, Fr. Guiana, 1856,

Sagot873(K).

17. Cnemidaria consimilis Stolze, sp. nov. Figure 22, Map 13.

Petiolus spinis usque ad 3 mm. longis instructus, petioli paleae abundantes,

albidae, vel stria centrali angustissima brunnea instructae; pinnae per unam

quartam vel mediam pattern distantiae inter apices pinnularum et costam incisae,

sinibus inter segmenta baud cartilagineis; pinnae laterales et terminales con-

similes; venae 1- vel 3-furcatae, venae infimae anastomosantes et areolas costales

efformantes; venae infimae basiscopicae e costulis plerumque egredientes; sori

uniseriati, inter costulam et marginem medii vel supramediani.

TYPE COLLECTION: Steyermark & Rabe 96260, Cerro de Rio Arriba,

Peninsula de Paria, Sucre, Venezuela, 9 Agosto 1966. HOLOTYPE: GH!

ISOTYPE: F!

Caudex not seen, reported to 1.5 m. long; leaves to 2.5 m. long and 1.2 m.

broad, terminating in a conform apical pinna; petiole to 1.5 m. long, muricate or

with spines to 3 mm. long, the scales abundant, lanceolate, whitish, or with a thin

brown to atropurpureus median stripe; rachis often spiny towards the base of the

lamina, non-alate, the scales sparse, pale to brownish, ovate to lanceolate, tri-

chomes lacking; pinnae sessile, cut one-fourth to almost halfway to the costa,

basal ones reduced and often deflexed; costae and costules with brown, amorphous

scales rare beneath or lacking, trichomes lacking; secondary segments subfalcate,

obtuse to rarely subapiculate, basal ones often crowding or overlapping the rachis;

veins once- to thrice-forked, basal ones commonly merging to form costal areoles,

basal basiscopic ones arising from the costule or its base; sori in a single line,

medial to supramedial between costule and segment margin; indusia yellowish-

or grayish-brown, more or less semicircular, 2- to 3-lobed, saucer-shaped to

deeply cup-shaped.

Although the central portion of the lamina of this species bears a

strong resemblance to that of Cnemidaria spectabilis, the petiole scales

and the shape of the apical section are quite distinct. Whereas the

lamina of C. spectabilis is abruptly reduced to a non-conform, broadly-

triangular apical section, the lamina of C. consimilis terminates in an

apical section similar in shape and size to one of the upper pinnae. The

difference in petiole scales is described fully in the discussion of C.

spectabilis.

FIG. 22. Cnemidaria consimilis, Steyermark & Rabe 96260 (GH, holotype).

67

68 FIELDIANA: BOTANY, VOLUME 37

C. consimilis might be confused also with C. speciosa, but the petiole

of the latter is smooth or rarely tuberculate, with scales clustered mostly

at the base and the brown median stripe quite broad. The petiole of

C. consimilis is quite spiny, or at least strongly muricate, and thickly be-

set throughout with scales that are either completely white or with a very

narrow brown median stripe, or a brown spot near the point of at-

tachment.

In forests, 600-1,200 m., Trinidad and the Peninsula de Paria, Vene-

zuela.

Additional specimens examined. — Venezuela: Sucre: Cerro de Humo,

Peninsula de Paria, Steyermark 94899 (F, GH, NY, U, VEN). Trinidad:

Mt. Tucuche, below the summit, Seifriz 4 (F). Forest, Mt. Tocuche,

N. L. Britton et al 1238 (GH, NY, US). Mt. Tucuche, Broadway

5296 (F), 7737 (US).

18. Cnemidaria speciosa Presl, Tent. Pterid. pi. 1, figs. 16, 17, 1836

(non Cyathea speciosa Willd. 1810). Figure 3c, Map 11.

TYPE COLLECTION: Poeppig 221 (Diar. 1144) Pampayaco, Peru, Juli 1829.

HOLOTYPE: PR or PRC. FRAGMENTS: NY! US! ISOTYPES: B! BR, P!

Hemitelia subincisa Kunze, Bot. Zeit. 2: 296. 1844. nom. nov. quoad Cnemi-

daria speciosa Presl, non Hemitelia speciosa (Willd.) Kaulf.

Hemistegia speciosa (Presl) Fee, Mem. Fam. Foug. 5 (Gen. Fil.): 351. 1850-52.

Cyathea subincisa (Kunze) Domin, Pteridophyta 264. 1929.

Caudex to 1 m. long and 3.5 cm. broad; leaves to 2.2 m. long and 0.7 m.

broad, terminating in a conform or subconform apical pinna; petiole to 1.2 m.

long, smooth or rarely tuberculate at the base, the scales lanceolate to ovate,

bicolorous, whitish with brown median stripe; rachis non-alate, scales and tri-

chomes lacking; pinnae sessile, broadest at or beyond the center, very deeply

crenate to lobed less than halfway to the costa, basal ones slightly, if at all, re-

duced and deflexed; costae and costules with dull brown, amorphous scales rare

beneath or lacking, trichomes lacking; secondary segments subfalcate, obtuse to

rarely subapiculate, basal ones crowding or overlapping the rachis; veins simple

to once-forked, basal ones commonly merging to form costal areoles, basal

basiscopic ones mostly arising from the costule or its base; sori in a single line

supramedial between costule and segment margin; indusia pale or yellowish, more

or less semicircular, saucer-shaped to deeply cup-shaped.

The lamina of Cnemidaria speciosa commonly terminates in a con-

form "apical pinna," the apical section closely resembling most of the

pinnae in shape and size. However, on some plants the apical section

is "subconform," i.e., it conforms to the shape of the pinnae except for

a pair of larger or elongate basal lobes. This is in marked contrast to

STOLZE: REVISION OF CNEMIDARIA 69

the "nonconform pinnatifid apex" of most species of Cnemidaria, which

are characterized by a very broad base and a triangular outline.

Cnemidaria speciosa resembles C. consimilis, especially in the central

portion of the lamina; however, in addition to the distinctions listed

under the discussion of the latter species, C. speciosa exhibits another

subtle difference. In most species of Cnemidaria (including C. consim-

ilis) the pinnae are generally broadest near the base, or at least below the

middle. C. speciosa is one of a few species having most pinnae slightly

broader at or beyond the middle. This may not always be evident upon

examination of a single pinna, but it becomes rather apparent on view-

ing the entire lamina.

Much confusion has existed between this species and Cyathea speciosa

Willd., and although there was a partial clarification by Kunze (1844)

and later by Maxon (1912), further elaboration is desirable especially

in reference to typification.

In 1836, Presl established the genus Cnemidaria, designating "Cnemi-

daria speciosa (Hemitelia speciosa Kaulf. nee Willd.)" and four other

taxa as the representatives of the new genus. Although he cited no

material for C. speciosa, he illustrated it by the figures 16 and 17, which

show the anastomosing of the lowest veins and serve to distinguish it

from Cyathea speciosa Willd. (earlier transferred to Hemitelia by Kaul-

fuss [1824]). Presl's publication made a new species, Cn. speciosa,

validated by the illustrations presented. The reference to "H. speciosa

Kaulf. nee Willd." was partly erroneous (it should have been "H. spe-

ciosa Kaulf. in herb.," not "Cyathea speciosa Willd."), and a fuller ex-

planation is to be found in Presl's (1847) later publication. Here the

citation of "Hemitelia speciosa Kaulf. herb.", of "Mart. ic. crypt, bras.

78 t. 48 f.II," and of "Kunze fil. Poepp. in Linn. IX 99 (excl. syn)" as

to specimens, provides the basis for Presl's action. In the same pub-

lication he restricts the genus to Cn. speciosa, removing the other four

species to Actinophlebia and Hemistegia. The Poeppig specimen at

Paris is labelled "No. 221. Hemitelia speciosa Kaulf.?," and it is this

identification (rather than Kaulfuss' description) that Presl argued

correctly did not agree with Cyathea speciosa Willd. Kunze's published

identification of the Poeppig collection was without question Hemitelia

speciosa Kaulf. [i.e. (Willd.) Kaulf.]. Maxon notes that "In the Presl

herbarium at Prague ... is a specimen of 'Cnemidaria speciosa' col-

lected in Peru by Poppig," and in the "Gefassbiindel" Presl cites a

specimen as "Pampayaco, Peru (Poeppig)." The fragment obtained by

Underwood from the Presl herbarium was from a Poeppig collection.

Finally, in the Syn. Fil., Poeppig cites "Diar. 1144" and places H.

70 FIELDIANA: BOTANY, VOLUME 37

speciosa Kaulf. as No. 263 in Kunze's enumeration. The Poeppig speci-

men at Paris is marked "No. 221 Hemitelia speciosa Klfs." (with the "1"

changed to a "4" in another handwriting and different ink) as well as

"Peru Poppig, Kunze 263." This specimen relates the Poeppig 221 to

Dira. 1144 and clearly establishes the fact that these are different num-

bers for the same collection.

In and at the edges of forests, along stream banks and mountain-

sides, 375-1, 900m., Peru, Bolivia.

Selected specimens examined. — Peru. Cuzco: San Lorenzo, Vargas

11714 (GH). Kosnipata-Pilcopata, Vargas 11294 (GH). Entre Atalaya

y Carbon, Prov. Paucartambo, Vargas 14653 (GH). Huanuco: Tingo

Maria, Asplund 12116 (US). Tingo Maria, roadside, Tryon & Tryon

5260 (F, GH, U). Junin: Near La Merced, Killip & Smith 23889,

(NY, US). Rio Paucartambo Valley, Killip & Smith 25289, (NY, US).

Pichis Trail, Yapas, Killip & Smith 25563 (GH, NY, US). Chancha-

mayo Valley, C. Schunke 52 (F). La Merced-Chanchamayo, Soukup

1069 (F). Bolivia. La Paz: Sobre el camino a Tipuani, Buchtien 4224

(F, MO, NY, US). Mapiri Region, San Carlos, Buchtien 42 & 43 (US).

Copacabana, Prov. Larecaja, Krukoff 11297 (F, GH, MO, NY, U, US).

Mapiri, Rusby 149 (GH, NY, US).

19. Cnemidaria ewanii (Alston) Tryon, Contr. Gray Herb. 200: 52.

1970. Figure 3d, Map 14.

Cyathea ewanii Alston, J. Wash. Acad. Sci. 48: 231, 1958. TYPE COLLEC-

TION: Ewan 16729, between Mocoa & Urcusique, Putumayo, Colombia, 9 Jan.

1945. HOLOTYPE: BM, ISOTYPES: GH! NO, US! PHOTO: F!

Caudex to 0.2 m. long and 3 cm. in diameter; leaves to 2.5 m. long and 0.5 m.

broad, terminating in a conform to subconform apical pinna; complete petiole

not seen, smooth to lightly muricate, the scales ovate to lanceolate, bicolorous,

whitish with brown median stripe; rachis non-alate, scales scattered, whitish

to bicolorous, trichomes lacking; pinnae sessile, mostly truncate at base, broadest

at or beyond the center, deeply crenate or rarely cut one-fourth to the costa,

basal ones scarcely reduced or deflexed; costae with whitish or pale amorphous

scales scattered beneath, trichomes lacking; costules with pale or dull brown scales

scattered or rare beneath, or lacking, trichomes lacking; secondary segments

or lobes obtuse to subapiculate; veins simple to once-forked, basal ones merging

to form costal areoles, basal basiscopic ones arising from the costule; sori in a

single line inframedial on the veins between costule and segment margin;

indusia pale to yellowish-brown, semicircular, subentire to several-lobed, saucer-

shaped.

Cnemidaria ewanii is one of the more advanced species of the genus,

as evidenced by the costal areoles, the shallow lobing of the pinnae,

and the lack of trichomes on the axes. It is very closely related to C.

STOLZE: REVISION OF CNEMIDARIA 71

speciosa, and it may be that the two merit only varietal distinction,

but until more complete collections are made of C. ewanii I prefer to

treat them as separate species. They are the only species in the genus

with pinnae broadest at or above the middle and share most other

characters as well. The pinnae of C. ewanii are more shallowly lobed,

with the upper ones often merely deeply crenate. The sori are inframedial

on the veins and the scales along the rachis and costae beneath are whit-

ish or pale in color. The sori of C. speciosa are supramedial and the

scales of the axes, when present, are dull brown.

Specimens of C. quitensis with less deeply dissected pinnae might

also be confused with C. ewanii, and the two species share the same

range. However, the pinnae of the former are slightly broader below

the center and there are spines on the petiole, and terete, recurved

trichomes on the rachis and costae beneath. The petiole of C. ewanii

is smooth or lightly muricate, and the axes lack trichomes.

In thickets and forests, 75-1,000 m., Colombia and Ecuador.

Additional specimens examined. — Colombia. Caqueta: Florencia,

Quebrada de las Perdices, Cuatrecasas 8854 (US). Sucre, Orillas del

Rio Hacha, Cuatrecasas 9022 (US). Sucre, Juzepczuk 6533 (US).

Choco: Istmina, on Rio San Juan, Killip 35454 (US). Putumayo: Um-

bria, alt. 325 m., Klug 1846 (F, GH, MO, NY, US). Orito, near road

at Rio Calderas, Plowman 2133 (COL, F, GH). Ecuador. "In sylvis f.

Bombonasa," Spruce 5365 (K, P). Guayas: Milagro, Crespi s.n. (US).

20. Cnemidaria roraimensis (Domin) Tryon, Contr. Gray Herb.

200: 52. 1970. Figure 23, Map 9.

Hemitelia roraimensis Domin, Kew Bull. 216, 1929. TYPE COLLECTION:

Appun 1127, Roraima, Br. Guiana 1863/64. HOLOTYPE: K! PHOTO: F!

Cyathea roraimensis Domin, Acta Bot. Bohem.9: 154. 1929.

Caudex not seen; leaves to ca. 1.2 m. long and 0.4 m. broad, terminating

abruptly in a nonconform, pinnatifid apical section; complete petiole not seen;

rachis non-alate, scales and trichomes lacking; pinnae sessile, cut one-fourth

to one-third to the costa; costae and costules lacking scales and trichomes; second-

ary segments or lobes rounded, obtuse to subapiculate, basal ones crowding or

overlapping the rachis; veins simple, basal ones merging to form costal areoles,

basal basiscopic ones arising from the costule; sori in a single line medial to supra-

medial on the veins between costule and segment margin, commonly forming a

V-shaped pattern on the segments; indusia brownish-yellow, semicircular, several-

lobed, saucer-shaped.

Cnemidaria roraimensis is known only from a small number of in-

complete specimens; future collections, hopefully, will include petioles

and sections of the caudex. Its affinities are with C. spectabilis, from

FlG. 23. Cnemidaria roraimensis, Appun 1127 (K, holotype).

72

STOLZE: REVISION OF CNEMIDARIA 73

which it differs in its more shallowly-lobed pinnae, simple veins, and

the V-shaped soral pattern on the segments (formed by the apical sori

lying close to the costule while the basal ones are much nearer the

margin). The pinnae of C. spectabilis are often cut nearly halfway to

the costa, the veins are usually branched at least once, and the sori

are uniformly arranged in a line supramedial between costule and seg-

ment margin.

Although probably not closely related, C. roraimensis and C. decur-

rens have at least a superficial resemblance, especially in their shal-

lowly-lobed pinnae. Besides the differences noted in the key, the two

are further distinguished by the relatively broad wing running well down

the sides of the rachis in most specimens of C. decurrens, whereas the

rachis is always non-alate in C. roraimensis. The shallow lobing of the

pinnae and the strongly areolate venation of both species place them

among the most highly advanced species in the genus.

Along stream banks and on mountainsides, 1,000-1,500 m., British

Guiana.

Selected specimens examined. — British Guiana: Quating Creek, 1864,

Appun 1035 (K). Mt. Ayanganna, Pakaraima Mts., Maguire et al.

40581 (NY). Roraima Range, 3,500 ft. alt., McConnell & Quelch

620 (K, NY).

21. Cnemidaria decurrens (Liebm.) Tryon, Contr. Gray Herb. 200:

52. 1970. Figure 24a, Map 12.

Hemitelia decurrens Liebm., Kongel Danske Vidensk. Selsk. Skr. V. 1: 286

(seors. 134), 1849. TYPE COLLECTION: Liebmann PL Mex. 2089 (No. 912),

Pr. Lobani, Distr. Chinantla, Dep. Oajaca, Mexico, 1842. HOLOTYPE: C!

FRAGMENT: US!

Hemitelia mexicana Liebm., torn. cit. 287 (seors. 135). TYPE COLLECTION:

Liebmann PL Mex. 2105 (3 sheets: Nos. 909, 910, 911), Pr. Lacoba, Dist. Chinan-

tla, Oajaca, Mexico, 1842. HOLOTYPE: C! FRAGMENT: US!

Hemistegia lucida Fee, Mem. Fam. Foug. 5 (Gen.Fil.): 351. 1850-52. TYPE

COLLECTION: Galeotti 6537, Prov. de Oaxaca, Mexico, 1842. HOLOTYPE:

BR, ISOTYPE: P! US!

Hemistegia elegantissima Fee, op. cit. 8: 110. 1857. TYPE COLLECTION:

Linden s.n., Mexico. Nothing of the type is known beyond Fee's reference, "Mex-

ique (Linden)," but according to the description, this plant can be nothing other

than Cnemidaria decurrens.

Hemistegia decurrens (Liebm.) Fourn., Mex. PI. 1: 135. 1872.

Hemistegia mexicana (Liebm.) Fourn. loc. cit.

Hemitelia guatemalensis Maxon, Contr. U.S. Natl. Herb. 16: 40. 1912. TYPE

COLLECTION: Salvin s.n., Alta Verapaz, Guatemala, 1800. HOLOTYPE: US.

ISOTYPE: GH! K! US!

Hemitelia lucida (Fee) Maxon, torn. cit. 39.

74 FIELDIANA: BOTANY, VOLUME 37

Cyathea guatemalensis (Maxon) Domin, Pteridophyta 264, 1929.

Cyathea liebmanii Domin, loc. cit. (nom. nov. for Hemitelia mexicana Liebm.,

non Cyathea mexicana S. & C. 1830.)

Cyathea lucida (Fee) Domin, loc.cit.

Cyathea decurrentiloba Domin, Acta Bot. Bohem. 9: 110. 1930. (nom. nov. for

Hemitelia decurrens Liebm., non Cyathea decurrens [Hook.] Copel. 1929.)

Cyathea elegantissima (Fee) Domin, torn. cit. 113.

Caudex rudimentary to 0.3 m. long; leaves to 2.5 m. long and 0.8 m. broad,

terminating in a gradually reduced apical section; petiole to 0.5 m. long, with

spines to 2 mm. long, the scales scattered to abundant, ovate to lanceolate, whitish

or bicolorous; rachis with membranous wing to 2 mm. wide running partially to

fully down each side or (rarely) lacking, the ovate to lanceolate scales whitish or

bicolorous, sparse to abundant beneath, the minute, terete, recurved trichomes

scattered to abundant beneath; pinnae subsessile, rounded to truncate at base,

subentire or crenate or cut one-third to the costa, basal ones scarcely or not at

all reduced and deflexed; costae with broad, amorphous, pale to whitish or bi-

colorous scales scattered beneath, trichomes lacking; costules with scales and

trichomes lacking; secondary segments or lobes (when present) obtuse to rarely

subapiculate; veins simple, basal ones commonly merging to form costal areoles,

basal basiscopic ones arising from the costule; sori in a single line medial

to inframedial on the veins between costule and segment margin; indusia pale

to yellowish-brown, semicircular to (rarely) almost fully circular, subentire to

several-lobed, saucer-shaped.

Cnemidaria decurrens has a variable leaf outline, probably correlated

with the age and size of the individual plant. Perhaps due to this vari-

ability, four distinct species had been previously recognized, most of

these originally represented by single collections. The types of Hemitelia

decurrens and H. mexicana represent the extremes: the pinnae of the

former nearly entire and the rachis strongly alate and copiously pubes-

cent and scaly. The type of H. mexicana has nearly entire upper pinnae,

but the large, central ones are lobed nearly one-third to the costa, a

wing is just barely evident along the rachis, and the pubescence and

scales sparse. However, examination of more recently collected ma-

terial shows these characters to be highly variable from one plant to

another, and hence there is little significant correlation between them.

Cnemidaria decurrens occurs at one border of the generic range, and,

as evidenced by the strongly areolate venation and the subentire to

shallowly lobed pinnae, is one of the most highly advanced species in

the genus. Its relationship is with C. choricarpa of Costa Rica, and was

probably derived from the latter's ancestral stock.

Hemitelia decurrens and H. mexicana were published at the same

time, so a choice of one of the names is necessary. Hemitelia decurrens

seems most appropriate because the pinnae are strongly decurrent,

and the rachis is commonly alate for most of its length. Furthermore,

STOLZE: REVISION OF CNEMIDAR1A 75

the majority of specimens more closely resemble the type of H. decur-

rens in that the margins of the pinnae are scarcely lobed. Leaves with

the pinnae lobed one-third to the costa as in the type of H. mexicana are

rather rare.

In the type folder of H. decurrens, at Copenhagen are two sheets, both

marked "Liebm. PI. Mex. 2089," each containing a small but com-

plete specimen. However, the labels of each are further designated with

different numbers, one "912," the other "913." The holotype is con-

sidered to be No. 912, and it is referred to by Fournier as such. The

other, numbered "913," is a paratype. In the Copenhagen type folder

of H. mexicana are three sheets, "Liebm. PI. Mex. 2105," and they

are further numbered "909," "910," and "911." These sheets are pre-

sumably from the same plant: obviously the apex, middle, and base of

one large lamina. Therefore, all three sheets must be considered parts

of the holotype of H. mexicana.

Forest slopes, 200-1,100 m., Mexico (Chiapas, Oaxaca) and Guate-

mala (Alta Verapaz).

Selected specimens examined. Mexico. Chiapas: Palenque, Munch 4

and s.n. (U.S. herb. no. 1791748} (US). Oaxaca: Dto. Choapam, Lo-

vani, Hallberg 1567 (F, NY). Tuxtepec, S. of Valle Nacional, Michel

5889, 5890 (NY). Tuxtepec, N. of Campamento Vista Hermosa,

Michel 5934 (NY). Guatemala: Alta Verapaz: Chapultepec Farm, Con-

treras 4822 (US). Finca Sepacuite, Cook & Griggs 107 (US). Finca

Seamay, Senehu, Hatch & Wilson 190 (GH, US). Finca Sepacuite,

Senehu, Wilson 21 8 A (F).

22. Cnemidaria karsteniana (Kl.) Tryon, Contr. Gray Herb. 200:

52, 1970. Figure 24b, Map 13.

Hemitelia karsteniana Kl., Allg. Gartenzeitung 20: 42. 1852. TYPE COLLEC-

TION: possibly Karsten 142, "Columbian, Decker 1851 am 31 Debr." HOLO-

TYPE: B! (?).

Cyathea karsteniana (Kl.) Domin. Pteridophyta 264, 1929.

Caudex to 0.6 m. long and 2.5 cm. in diameter; leaves to 2.5 m. long and 0.8

m. broad, terminating in a conform or subconform apical pinna (often with a pair

of enlarged basal lobes); petiole to 1 m. long, muricate to short-spiny, the scales

lanceolate to linear-lanceolate, bicolorous, brown with narrow, whitish margins;

rachis non-alate, lacking scales and trichomes; pinnae subsessile, crenate or

shallowly lobed, truncate at base, the tissue sometimes crowding or overlapping

the rachis; costae and costules with scales and trichomes lacking; veins mostly

once-forked, basal ones merging to form costal areoles, basal basiscopic ones

arising from the costule; sori in a single line mostly inframedial on the veins

between costule and segment margin; indusia yellowish-brown, more or less semi-

circular, subentire to lobed, saucer-shaped.

76

FIELDIANA: BOTANY, VOLUME 37

This and Cnemfdaria nervosa are the most highly advanced species

in the genus. Both have strongly areolate venation along the costae, sub-

entire to shallowly lobed pinnae, and are completely lacking in scales

FIG. 24. Portions of pinnae, showing venation; simple, a, Cnemidaria decur-

rens, Hatch & Wilson 190 (GH), and, c, C. nervosa, Mexia 6291 (GH, isotype);

once-forked, b, C. karsteniana, Tschudi 99 (VEN).

or trichomes except on the petiole. They are not, however, closely re-

lated and can easily be distinguished by the characters noted in the key.

They have apparently arisen along different branches of the phyletic line

and have attained their similar advanced characteristics through par-

allel evolution. It is likely that C. nervosa stems from the same line as

C. speciosa and C. ewanii, whereas C. karsteniana is more closely re-

lated to C. consimilis.

There is a specimen of Hemitelia karsteniana at Kew, designated as

a type, with the label data simply "Hort. Lips. Mett., Herb. Hooker-

iana," but I find no evidence to indicate this is type material. However,

at Berlin, a specimen of Karsten 142, marked "Hemitelia (Cnemidaria)

karsteniana Klotzsch," bears a handwritten label nearly identical with

that of Karsten 143 which is determined as "Hemitelia klotzschiana"

(see also discussion under C. spectabilis var. spectabilis) . Both of these

were collected in 1851, and both species were described together a

year later by Klotzsch, rather strong evidence that these were the speci-

mens on which he had based his original descriptions.

Slopes of cloud forests, 1,000-1,600 m. Apparently confined to

northern Venezuela (Aragua, Carabobo, Sucre). Karsten 142, cited

above, bears no collection locale other than "Columbien," a designation

Karsten used for Venezuelan as well as Colombian collections.

Selected specimens examined. — Venezuela. Aragua: Alto de Choroni,

Chardon 188 (US, VEN). Highway between Maracay & Choroni,

STOLZE: REVISION OF CNEMIDARIA 77

Lasser 210 (GH, US, YEN). Cloud forests of Rancho Grande, Pittier

13982 (F, US, YEN). Maracay, 1939, Vogl s.n. (F). Parque Nacional,

LI Williams 10758, 10786 (F). Carabobo: Funk & Schlim 613 (F).

Sucre: Cerro Patao, Penin. de Paria, Steyermark & Agostini 91093 (US,

YEN).

23. Cnemidaria nervosa (Maxon) Tryon, Contr. Gray Herb. 200:

52, 1970. Figure 24c, Map 14.

Hemitelia nervosa Maxon, J. Wash. Acad. Sci. 34: 309, 1944. TYPE COLLEC-

TION: Mexia 6291, mouth of Rio Santiago, Loreto, Peru, Dec. 18, 1931. HOLO-

TYPE: (3 sheets) US! ISOTYPES: F! GH! NY!

Caudex not seen, reported to be less than 1 m. long; leaves to 2.5 m. long and

0.7 m. broad, terminating in a conform or subconform apical pinna; petiole to 1

m. long, smooth or- muricate, the scales mostly near the base, lanceolate to lin-

ear-lanceolate, bicolorous, brown with narrow whitish margins; rachis non-alate,

lacking scales and trichomes; pinnae short-stalked, subentire to broadly serrate,

rounded to broadly cuneate at- base; costae and costules with scales and trichomes

lacking; veins simple, basal ones merging to form costal areoles, basal basiscopic

ones arising from the costule; sori in a single line inframedial on the veins be-

tween costule and margin; indusia pale or yellowish, commonly circular, com-

pletely surrounding the receptacle, entire to lobed, flattened or saucer-shaped.

This is one of the most distinctive species in the genus. Its pinnae

are often nearly entire, with rounded bases, and commonly reach a

breadth of 9 cm. Probably the most highly advanced species of Cnemi-

daria, it could perhaps be confused only with C. karsteniana. (A discus-

sion of both is included under the latter. )

One of the unique characters of C. nervosa is the completely circular

indusium, which fully surrounds the receptacle. It is flat or slightly con-

cave and looks very much like a saucer. Only C. cocleana shares this

feature. Other species of Cnemidaria have more or less semicircular in-

dusia, positioned on the costular side of the receptacle.

Rain forests, 300-450 m., Peru (Amazonas, Loreto) and Ecuador

(Santiago-Zamora) .

Additional specimens examined. — Peru. Amazonas: Valley of Rio

Maranon, Prov. de Bagua, Wurdack 2059 (NY, US). Ecuador: San-

tiago-Zamora: Taisha, banks of Rio Guaguayme, Cazalet & Penning-

ton 7746 (NY).

DUBIOUS NAMES

HEMISTEGIA AMERISTONEURA Fee, Mem. Fam. Foug. VIII: 110.

1857. TYPE COLLECTION: Poiteau s.n., Guayane Francaise.

Hemitelia ameristoneura (Fee) C. Chr., Ind. Fil. 347. 1905.

FIG. 25. Cnemidaria spores: a, C. chiricana, proximal view (1260 X); b, C.

chiricana, fine relief (6300 X); c, C. mutica var. mutica, proximal view (1350

X); d, C. choricarpa, proximal view (1200 X); e, C. uleana var. uleana, proxi-

mal view (1300 X); f, C. ewanii, distal view (1250 X).

78

FIG. 26. Cnemidaria spores: a, C. horrida, lateral view (1250 X); b, C.

horrida, distal view (1270X); c, C. grandi folia var. grandi folia, proximal view

(1250 X); d, C. spectabilis var. spectabilis, proximal view (1220 X); e, C. decur-

rens, proximal-lateral view (1300X); f, C. karsteniana, proximal-lateral view

(1270 X).

79

80 FIELDIANA: BOTANY, VOLUME 37

Cyathea ameristoneura (Fee) Domin, Pteridophyta 263. 1929.

I have not seen the type specimen, and from the description it is

difficult to determine with certainty if this is a distinct species or one

of the known species of Cnemidaria. It could be either C. cruciata or

C. spectabilis, the two representatives of the genus known in French

Guiana. The only Poiteau specimens I have seen are those of C. cruciata,

and Fee's description perhaps matches this species more closely than it

does C. spectabilis: ". . .longeur des frondules 14-17 centim., sur 3

centim. de largeur." Rarely do larger pinnae of C. spectabilis measure

less than 4.5 cm. wide. However, too little evidence is currently avail-

able to place this name with certainty.

HEMITELIA MONILIFERA J.Sm., London J. Bot. 1: 662. 1848. nom.

nud.

Hemistegia monilifera Presl, Gefassb. Stipes der Farm 47. 1847 (preprint of

Abh. Bohm Ges. Wis. V. 5: 355. 1848). nom. nud.

The name Hemitelia monilifera first appears in Smith's "Genera of

Ferns," and later is listed by Presl (1847) under Hemistegia. I have

seen no authentic specimens, therefore I am uncertain whether the

plants thus referred are actually Cnemidaria. Neither Smith nor Presl

included descriptions of any kind, although it may be inferred from the

former's general remarks that H. monilifera belongs to the ". . .true

Hemiteliae of Mr. Brown," in which "the lower venules anastomose."

Nothing further may be positively determined at this time.

EXCLUDED SPECIES

On the basis of the evidence presented in this revision, five species

included in Cnemidaria by Tryon (1970) must now be excluded. These

are herewith placed in Cyathea, with new combinations where necessary:

CYATHEA conformis (Tryon) Stolze, comb. nov.

Hemitelia conformis Tryon, Rhodora 62: 1. 1960.

Cnemidaria conformis (Tryon) Tryon, Contr. Gray Herb. 200: 51. 1970.

CYATHEA dissimilis (Morton) Stolze, comb. nov.

Hemitelia dissimilis Morton, Fieldiana Bot. 28: 8. 1951.

Cnemidaria dissimilis (Morton) Tryon, Contr. Gray Herb. 200: 52. 1970.

CYATHEA INTEGRIFOLIA (Kl.) Domin, Pteridophyta 264. 1929.

Hemitelia integrifolia Kl., Linnaea 18: 539. 1844.

Cnemidaria integrifolia (Kl.) Tryon, Contr. Gray Herb. 200: 52. 1970.

CYATHEA PANAMENSIS Domin, Pteridophyta 264. 1929.

Hemitelia marginalis J. Sm. 1842. nom. nud. (non Cyathea marginalis Domin

1929.)

STOLZE: REVISION OF CNEMIDARIA 81

Hemitelia petiolata Hook., Spec. Fil. I. 31, /. 16. 1844. (non Cyathea peti-

olata J. Sm. 1841).

Hemistegia marginalis Presl, Gefassb. Stipes der Farm, 47. 1847 (preprint of

Abh. Bohm Ges. Wiss V. 5: 355. 1848).

Cnemidaria petiolata (Hook.) Copel., Gen. Fil. 97. 1947.

CYATHEA SPECIOSA Willd., Spec. PI. V: 490. 1810 (non Cnemidaria

speciosa Presl 1836).

Hemitelia speciosa (Willd.) Kaulf., Enum. Fil. 252. 1824.

Hemitelia lindenii Hook., Icon. PI. /. 706 1848.

Cnemidaria lindenii (Hook.) Tryon, Contr. Gray Herb. 200: 52. 1970.

Three other species have occasionally been confused with Cnemidaria,

either because of the anastomosing of some veins, or because the lamina

is not highly dissected. These are also excluded from Cnemidaria and

new combinations are herewith provided as needed :

CYATHEA hombersleyii (Maxon) Stolze, comb. nov.

Hemitelia hombersleyi Maxon, J. Wash. Acad. Sci. 25: 528. 1935.

CYATHEA WILSONII (Hook.) Domin, Pteridophyta 264. 1929.

Hemitelia wilsonii Hook, in Hook. & Bak. Syn. Fil. 30. 1865.

CYATHEA woronovii (Maxon & Morton) Stolze, comb. nov.

Hemitelia woronovii Maxon & Morton, Amer. Fern J. 36: 91. 1946.

One other species which bears superficial resemblance to species of

Cnemidaria was described by Kuhn (1869) as:

HEMITELIA VENOSA Reichb. Herb. Linnaea 36:161. 1869.

Unfortunately, the only material I have been able to locate is a type

fragment at U.S. National Herbarium, consisting of a single pair of seg-

ments, and a photograph taken by Drs. Rolla and Alice Tryon of the

type collection at Berlin. Little can be determined from the frag-

ment except that the veins are free and somewhat connivent to the sinus,

and that the indusia are hemitelioid. The type photograph reveals an

unnumbered specimen of Appun, from "Caracas," consisting of what

appears to be a rachis with three fragmentary pairs of pinnae. The

pinnae bear a few subfalcate, obtuse segments, cut fully (or nearly) to

the costa, which is a condition atypical of Cnemidaria. Kuhn's descrip-

tion sheds no further light on the plant's generic status. It could there-

fore be a strange species of Cnemidaria, or, more probably, a Cyathea.

It is of little use to conjecture further.

82

MAPS 8-9. 8, Cnemidaria singularis, dot; C. uleana var. uleana, square; C.

uleana var. abitagnensis, diamond. 9, C. spectabilis var. spectabilis, dot; C.

spectabilis var. colombiensis, triangle; C. cruciata, square; C. roraimensis, dia-

mond.

83

MAPS 10-11. 10, Cnemidaria horrida, diamond; C. grandifolia var. gran-

difolia, circle; C. grandifolia var. obtusa, open square. 11, C. speciosa, square;

C. alatissima, dot; C. quitensis, diamond.

84

MAPS 12-14. 12, Cnemidaria chocoensis, square; C. choricarpa, dot; C.

cocleana, diamond; C. decurrens, triangle. 13, C. consimilis, square; C. kar-

steniana, dot. 14, C. nervosa, dot; C. ewanii, diamond.

85

REFERENCES1

CHRISTENSEN, C.

1905. Index Filicum, Hafniae 1906. Suppl. 1906-1912: Hafniae 1913. Suppl.

1913-1916: Hafniae 1917.

COPELAND, E. B.

1947. Genera Filicum. Chronica Botanica. Waltham, Massachusetts.

DOMIN, K.

1929. New ferns from Tropical America and the West Indies. Kew Bull. 7:

215-222.

1929. Pteridophyta. Praha.

1929. The Pteridophyta of the Island of Dominica. Mem. Roy. Czech Soc.

Sci. 2.

1930. The species of the genus Cyathea J. E. Smith. Acta Bot. Bohem. 9:

85-174.

ERDTMAN, G. and P. SORSA

1971. Pollen and spore morphology /plant taxonomy: Pteridophyta. Almqvist

and Wiksell, Stockholm, Sweden.

FEE, A. L. A.

1852. Cinquieme Memoire: Genera Filicum exposition des genres de la famille

des Polypodiacees (Mem. Fam. Foug. 5). Paris and Strasbourg.

FOURNIER, E.

1872. Mexicanas Plantas. ... 1: Cryptogamia. Parisiis.

GRAHAM, A. and D. M. JARZEN

1969. Studies in neotropical paleobotany, 1: The Oligocene communities of

Puerto Rico. Ann. Missouri Bot. Gard. 56: 308-357.

HANNEN, T. VAN DER and E. GONZALEZ

1960. Upper Pleistocene and Holocene climate and vegetation of the "Sabana

de Bogota." Leidse Geol. Meded. 25: 261-315.

HOLLTUM, R. E. and U. SEN

1961. Morphology and classification of the tree ferns. Morphology 11 (4):

406-420.

HOOKER, W. J. and J. G. BAKER

1865-1868. Synopsis Filicum, ed. I. London.

KAULFUSS, G. F.

1824. Enttmeratio Filicum. Lipsiae.

KLOTZSCH, J. F.

1852. Einige neue Baumfarrn. Allg. Gartenzeitung 20: 41-49.

KUHN, M.

1869. Reliquiae Mettenianae. Linnaea 36:41-169.

1A11 abbreviations of periodical publications according to Botanico-Periodi-

cum-Huntianum, Hunt Botanical Library, Pittsburgh, Pennsylvania, 1968.

86

STOLZE: REVISION OF CNEMIDARIA 87

KUNZE. G.

1844 Hooker, Species Filicum . . . Hot. Zeit. (Berlin) 2: 294-299.

LANJOUW, J. and F. A. STAFLEU

1964. Index Herbariorum. Reg. Veget. 31: 1-251.

LIEBMANN, F.

1849. Mexicos Bregner, en systematisk, critisk, plantegeographisk, Under-

sogelse. (Repaged reprint of Kongel Danske Vidensk. Selsk, Skr. 1848).

MAXON, W. R.

1912. Studies of tropical American ferns, No. 3. Contr. U. S. Natl. Herb. 16:

25-49.

1914. A new species of Hemitelio, Section Cnemidaria, from Panama. Contr.

U. S. Natl. Herb. 17 (4): 413-414.

PRESL, C. B.

1836. Tentamen Pteridographiae, VIII: Cnemidaria. 56-57. Praha.

1847. Die Gefassbiindel im Stipes der Farm: Cyatheaceae. 38-48. (Preprint

of Abh. Bohm Ges. Wiss. V, 5: 346-356. 1848.)

SCHLECHTENDAL, D. F. L. VOH

1856. Ueber Poly podium horridum L. Eine kritische betrachtung. Bot. Zeit.

(Berlin) 14: 465-475.

SMITH, J.

1841. An arrangement and definition of the genera of ferns. London. J. Bot.

1: 659-668.

TRYON, R. M.

1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200: 1-53.

WILLDENOW, K. L.

1810. Caroli a Linne Species Plantarum, ed. quarta, Vol. V. Berolini.

INDEX TO COLLECTORS' NUMBERS

ABBOTT, W. L.

349 horrida

472 horrida

1372 horrida

1618 horrida

2545 horrida

2651 horrida

2726 horrida

ACOSTA-SOLIS, M.

12720 quitensis

AGUILAR, P.

246 speciosa

ALLARD, H. A.

13599 horrida

13794 horrida

14220 horrida

14303 horrida

2 1 605 speciosa

ALLORGE, P.

s.n. (U) grandi folia var. grandi-

folia

ANDRE, E.

1052 horrida

APOLLINAIRE-MARIE, BRO.

12 horrida

APPUN, C. F.

1035 roraimensis

1127 roraimensis

ARCHER, W. A.

1764 spectabilis var. colombiensis

ASPLUND, E.

9230 horrida

12116 speciosa

AUGUSTO, BRO.

859 horrida

BAILEY, L. H.

758 grandi folia var. grandi folia

761 grandifolia var. grandifolia

781 grandifolia var. grandifolia

BAILEY, L. H. and E. Z. BAILEY

80 grandifolia var. grandifolia

274 grandifolia var. grandifolia

651 horrida

T-25 spectabilis var. spectabilis

BALCH, A.

s.n. (NY) horrida

BEARD, P.

1 194 grandifolia var. obtusa

1434 grandifolia var. obtusa

BERNARDI, A. L.

836 spectabilis var. spectabilis

BLOMQUIST, H. L.

11684 horrida

BOLDINGH, I.

222 IB grandifolia var. grandifolia

2225B grandifolia var. grandifolia

Box, H. E.

283 grandifolia var. grandifolia

490 grandifolia var. grandifolia

1963 grandifolia var. obtusa

BRADE, A. C.

102 mutica var. mutica

451 mutica var. grandis

6668a uleana var. uleana

6901 uleana var. uleana

9252 uleana var. uleana

10325 uleana var. uleana

BRENES, A. M.

4992 mutica var. contigua

22003 mutica var. mutica

BRITTON, E. G.

588 horrida

BRITTON, E. G. et al.

433 spectabilis var. spectabilis

551 horrida

BRITTON, N. L.

3986 horrida

BRITTON, N. L. et al.

510 grandifolia var. grandifolia

551 horrida

831 horrida

1011 spectabilis var. spectabilis

1238 consimilis

1748 spectabilis var. spectabilis

2075 horrida

2217 horrida

3897 horrida

5187 horrida

6356 horrida

6386 horrida

6576 horrida

8401 horrida

BROADWAY, W. E.

567 spectabilis var. spectabilis

3886 spectabilis var. spectabilis

4540 spectabilis var. spectabilis

4720 grandifolia var. obtusa

4850 spectabilis var. spectabilis

5296 consimilis

5379 spectabilis var. spectabilis

7137 consimilis

s.n. (1904) grandifolia var. obtusa

s.n. (1905) grandifolia var. obtusa

88

INDEX TO COLLECTORS' NUMBERS

89

BUCHTIEN, O.

42 speciosa

43 speciosa

1047 speciosa

1087 speciosa

5223 speciosa

5224 speciosa

BURGER, W. C.

3950 mutica var. mutica

BURGER, W. C. and R. L. LIESNER

6771 mutica var. mutica

7135 choricarpa

7262 choricarpa

BURGER, W. C. and G. MATTA

4756 choricarpa

BURGER, W. C. and R. G. STOLZE

5270 mutica var. grandis

5328 mutica var. mutica

5499 choricarpa

5685 mutica var. mutica

5936 mutica var. mutica

6096 mutica var. mutica

CAMPBELL, W.

s.n. (1958) grandifolia var. grandi-

folia

CARDENAS, M.

1263 speciosa

CAZALET, P. and T. PENNINGTON

7711 horrida

7746 nervosa

CHAMBERS, K. L.

2765 grandifolia var. grandifolia

CHARDON, C. E.

188 karsteniana

CHRYSLER, M. A.

1803 horrida

CHRYSLER, M. A. and W. E. ROEVER

5443 mutica var. grandis

CLEMENT, Bro.

5235 horrida

CLUTE, W. N.

266 horrida

CONTRERAS, E.

4822 decurrens

COOK, O. F. and R. F. GRIGGS

107 decurrens

COOLEY, G. R.

8240 grandifolia var. obtusa

8377 grandifolia var. obtusa

8419 grandifolia var. obtusa

8421 grandifolia var. obtusa

8781 grandifolia var. grandifolia

COOPER, J. J.

s.n. (US) mutica var. grandis

CORNMAN, L. R.

896 mutica var. mutica

1058 mutica var. mutica

1285 mutica var. mutica

COWAN, R. S.

38982 spectabilis var. spectabilis

COWELL, J. F.

581 horrida

CRESPI, C.

s.n. (US) ewanii

s.n. (US) horrida

CROSBY, M. R. and W. R. ANDERSON

1042 horrida

CUATRECASAS, J.

4552 horrida

8640 uleana var. abitaguensis

8854 ewanii

9022 eivfl/m

CURRAN, H. M. and M. HAMAN

676 grandifolia var. obtusa

683 grandifolia var. obtusa

736 grandifolia var. obtusa

DANIEL, Bro.

913 horrida

1897 horrida

DAY, E. H.

137 horrida

336 spectabilis var. spectabilis

591 horrida

DELGADO, E.

81 horrida

DUDLEY, T. R.

10459 uleana var. uleana

11312 uleana var. uleana

13281 alatissima

13282 alatissima

DUGAND, A. and R. JARAMILLO

3813 horrida

DUSEN, P.

2109 uleana var. uleana

14245 uleana var. uleana

17306 uleana var. uleana

18124 uleana var. uleana

Duss, Pere

1 607 grandifolia var. grandifolia

1610 grandifolia var. grandifolia

4151 grandifolia var. grandifolia

4153 grandifolia var. grandifolia

4155 grandifolia var. grandifolia

4434 grandifolia var. grandifolia

4435 grandifolia var. grandifolia

4449 grandifolia var. grandifolia

4451 grandifolia var. grandifolia

4452 grandifolia var. grandifolia

4605 grandifolia var. grandifolia

EGGERS, H. F.

867 grandifolia var. grandifolia

1423 spectabilis var. spectabilis

2738 horrida

5030 horrida

5859 grandifolia var. obtusa

6017 grandifolia var. obtusa

6035 grandifolia var. obtusa

673 1 grandifolia var. obtusa

6804 grandifolia var. obtusa

EKMAN, E. L.

2875 horrida

3771 horrida

10668 horrida

1154Q horrida

90

INDEX TO COLLECTORS' NUMBERS

EKMAN, E. L. (continued)

H-2875 horrida

H-4683 horrida

ELLIOTT, W. R.

s.n. (1887) grandifolia var. obtusa

EWAN, J. A.

15669 horrida

16729 ewanii

16790 horrida

16812 quitensis

17084 grandifolia var. grandifolia

FELDMAN, J.

s.n. (IJ) grandifolia var. grandi-

folia

FENDLER, A.

25 spectabilis var. spectabilis

385 horrida

386 karsteniana

480 spectabilis var. spectabilis

FERREYRA, R.

1921 speciosa

FISHER, G. L.

67 horrida

FLEMING, H. and F.

47 spectabilis var. spectabilis

FREDHOLM, A.

3340 horrida

FRIEND, E. A.

77 grandifolia var. grandifolia

FUERTES, M.

769 horrida

917B horrida

1552 horrida

FUNCK*

769 karsteniana

FUNK, N. and L. SCHLIM

613 karsteniana

GALEOTTI, H.

6537 decurrens

GARBER, A. P.

133 horrida

GARCIA, J.

45 karsteniana

GARCIA-BARRIGA, H.

10734 horrida

13120 quitensis

GASTONY, G.

7 horrida

8 horrida

39 horrida

41 horrida

GASTONY, G. and S.

756 mutica var. mutica

764 mutica var. mutica

781 mutica var. mutica

GASTONY, G. et al.

655 horrida

704 horrida

GINES, Bro.

148 horrida

2621 grandifolia var. obtusa

3434 grandifolia var. obtusa

3451 grandifolia var. obtusa

3462 grandifolia var. obtusa

GLAZIOU, A.

2420 uleana var. uleana

GOMEZ, L. D.

2246 mutica var. contigua

GONZALEZ, A.

653 horrida

GRANT, M. L.

9126 horrida

GRUBB, P. J. et al.

1238 horrida

1392 horrida

GUEVARA-AMORTEGUI, B.

289 horrida

HAHN, L.

52 grandifolia var. grandifolia

HALLBERG, B.

1557 apiculata

1567 decurrens

HARRIS, W. and N. L. BRITTON

10697 horrida

10709 horrida

HART, J. H.

174 horrida

HATCH, W. R. and C. L. WILSON

1 90 decurrens

HEKKING, W. H. A.

1421 spectabilis var. spectabilis

1435 spectabilis var. spectabilis

1441 spectabilis var. spectabilis

HELLER, A. A.

1043 horrida

HENRI-STANISLAUS, Bro.

1644 horrida

HIORAM, Bro.

277 horrida

HIORAM, Bro. and Bro. CLEMENT

6526 horrida

HIORAM, Bro. and C. MAUREL

2473 horrida

4695 horrida

HODGE, W. H.

8 grandifolia var. grandifolia

9 grandifolia var. grandifolia

10 grandifolia var. grandifolia

1332 grandifolia var. grandifolia

1711 grandifolia var. grandifolia

2378 grandifolia var. grandifolia

3438 grandifolia var. grandifolia

HODGE, W. H. and B. T.

1814 grandifolia var. grandifolia

HOLDRIDGE, L. R.

2208 horrida

"'Indicates collectors for whom I was unable to locate initials or first names.

INDEX TO COLLECTORS' NUMBERS

91

HOLM, T.

169a horrida

HOMBERSLEY, A.

30 spectabilis var. spectabilis

210 spectabilis var. spectabilis

HOWARD, R. A.

5222 horrida

5376 horrida

6427 horrida

11964 grandi folia var. grandi 'folia

HOWARD, R. A. and L. I. NEVLING

15585 horrida

15763 horrida

HUMPHREY, J. E.

3340 horrida

HUNNEWELL, F. W.

11530/zomWo

HUSNOT, T.

388 grandifolia var. grandifolia

IDROBO, J. and R. SCHULTES

735 horrida

IMRAY *

14 grandifolia var. grandifolia

JACK, J. G.

6438 horrida

6477 horrida

7341 horrida

JENMAN, G. S.

1 24 spectabilis var. spectabilis

s.n. (1895) spectabilis var. specta-

bilis

s.n. (1899) spectabilis var. specta-

bilis

JIMENEZ, A.

1644 mutica var. mutica

2015 mutica var. mutica

JIMENEZ, J.

3069 horrida

JOHNSTON, J. R.

191 grandifolia var. obtusa (pro

parte)

191 spectabilis var. spectabilis (pro

parte)

JONES, G. C. and D. H. NORRIS

1187 horrida

JONES, G. N.

10993 horrida

JURGENSEN, C.

"273" apiculata

873 apiculata

905 decurrens

JUZEP'CZUK, S.

6510 horrida

6533 ewanii

KAPPLER, A.

1771 spectabilis var. spectabilis

KARSTEN, H.

142 karsteniana

143 spectabilis var. spectabilis

s.n. (B) karsteniana

s.n. (F) karsteniana

KARWINSKY, W. F.

s.n. (F) decurrens

KILLIP, E. P.

5039 spectabilis var. colombiemis

5156 mutica var. mutica

5162 mutica var. mutica

523 1 mutica var. grandis

5248 mutica var. grandis

5294 mutica var. chiricana

5297 mutica var. chiricana

5347 mutica var. chiricana

5350 mutica var. chiricana

5391 mutica var. grandis

7774 quitensis

34434 horrida

35377 spectabilis var. colombiensis

35454 ewanii

KTLLFP, E. P. and T. E. HAZEN

10159/zormfo

KILLIP, E. P. and A. C. SMITH

23889 speciosa

24536 speciosa

24539 speciosa

24548 speciosa

25289 speciosa

25563 speciosa

25737 speciosa

26341 speciosa

26574 speciosa

KLUG, G.

1846 ewanii

3182 horrida

KRAMER, K. U. and W. H. A.

HEKKING

3 195 spectabilis var. spectabilis

KRUKOFF, B. A.

1 1297 speciosa

KUNKEL, G.

596 speciosa

KUNTZE, O.

s.n. (NY) spectabilis var. specta-

bilis

s.n. (US) horrida

LANKESTER, C. H.

726 mutica var. mutica

732 mutica var. mutica

LASSER, T.

210 karsteniana

2216 karsteniana

LASSER, T. and L. ARISTIGUIETA

3413 grandifolia var. obtusa

LAVASTRE, B. A.

1887 horrida

1889 horrida

LECHLER, W.

2172 speciosa

s.n. (F) speciosa

LEHMANN, F. C.

7367 horrida

LELLINGER, D. B.

619 grandifolia var. grandifolia

92

INDEX TO COLLECTORS' NUMBERS

LELLINGER, D. B. and

E. R. de la SOTA

277 choricarpa

43 1 spectabilis var. colombiensis

728 quitensis

763 chocoensis

895 chocoensis

LENT, R. W.

28 mutica var. grandis

LEON, Bro. and C. MAUREL

3823 horrida

LEONARD, E. C.

7863 horrida

8209 horrida

9189 horrida

9191 horrida

9193 horrida

9373 horrida

LEONARD, E. C. and G. M.

12275 horrida

14238 horrida

14312 horrida

LEPRIEUR, M.

s.n. (K) spectabilis var. spectabilis

LIEBMANN, F.

909 (PI. Mex. 2105) decurrens

910 (PI. Mex. 2105) decurrens

911 (PI. Mex. 2105) decurrens

912 (PI. Mex. 2089) decurrens

913 (PI. Mex. 2089) decurrens

s.n. (US) decurrens

LINDEMAN, J. C.

6124 spectabilis var. spectabilis

LINDEN, J. J.

448 horrida

1738 horrida

LLOYD, F. E.

263 grandifolia var. grandifolia

LOPEZ, M.

143 horrida

LORD, M. L.

s.n. (1952) grandifolia var. grandi-

folia

LUERSSEN, C.

s.n. (P) ("Hort. Bot. Lips.") bella

LUTZ, B.

727 uleana var. uleana

729 uleana var. uleana

MAGDEFRAU, K.

6 horrida

MAGUIRE, B. et al.

40581 roraimensis

MAXON, W. R.

307 mutica var. grandis

382 mutica var. mutica

434 mutica var. mutica

451 mutica var. mutica

453 mutica var. mutica

523 mutica var. contigua

869 horrida

2387 horrida

2456 horrida

4034

4104

4187

4336

5474

5517

5519

5521

5682

8879

9346

horrida

mutica var.

horrida

chiricana

grandis

MAXON, W. R. and A. D. HARVEY

8023 mutica var. grandis

8032 mutica var. mutica

8042 mutica var. grandis

8066 mutica var. mutica

8136 mutica var. contigua

8220 mutica var. contigua

MAXON, W. R. and E. P. KILLIP

12 horrida

779 horrida

1550 horrida

MCCONNELL, F. V. and J. J. QUELCH

620 roraimensis

METCALF, R. D. and J. CUATRECASAS

30122 tryoniana

MEXIA, Y.

6291 nervosa

7320 horrida

8479 quitensis

MlCKEL, J. T.

994 apiculata

5676 apiculata

5717 apiculata

5889 decurrens

5890 decurrens

5934 decurrens

5935 apiculata

MILLER, G. S.

1180 horrida

s.n. (1924) grandifolia var. obtusa

MILLER, O. O. and J. R. JOHNSTON

164 spectabilis var. spectabilis

MOLINA, A. et al.

17954 choricarpa

MORITZ, J.

290 horrida

MORTON, C. V.

4240 horrida

4326 horrida

5072 grandifolia var. obtusa

5207 grandifolia var. obtusa

5717 grandifolia var. obtusa

MUNCH, G.

4 decurrens

s.n. (US) decurrens

MURILLO, M. T.

70 horrida

NASH, G. V.

239 horrida

INDEX TO COLLECTORS' NUMBERS

93

NlSMAN, C.

7 mutica var. grandis

14 mutica var. mutica

15 mutica var. mutica

22 mutica var. mutica

35 choricarpa

62 mutica var. mutica

87 choricarpa

93 mutica var. grandis

123 mutica var. grandis

127 mutica var. mutica

146 choricarpa

155 mutica var. mutica

173 choricarpa

OWNBEY, M.

2707 horrida

PALMER, C.

64 mutica var. mutica

PENNELL, F. W.

1577 horrida

PEREIRA, E.

312 uleana var. uleana

PHILIPSON, W. R. et al.

1658 horrida

PlTTIER, H.

1679 horrida

1837 mutica var. mulica

4835 choricarpa

5945 spectabilis var. spectabilis

7091 spectabilis var. spectabilis

7794 spectabilis var. spectabilis

10966 choricarpa

10969 p.p. mutica var. mutica

10969 p.p. choricarpa

11510 horrida

13906 karsteniana

13982 karsteniana

PLOWMAN, T.

2133 ewanii

POEPPIG, E. F.

"221" speciosa

224 speciosa

POITEAU, A.

139 cruciata

s.n. (1824) cruciata

POLLARD, C. L. and W. PALMER

118 horrida

154 horrida

PORTER, T. C.

s.n. (US) &<?//«

PROCTOR, G. R.

3894 horrida

3911 horrida

5117 horrida

16773 grandifolia var. grandifolia

16904 grandifolia var. obtusa

17523 grandifolia var. .grandifolia

18092 grandifolia var. grandifolia

1 9097 grandifolia var. grandifolia

19318 grandifolia var. grandifolia

19501 grandifolia var. grandifolia

20379 grandifolia var. grandifolia

21553 spectabilis var. spectabilis

21699 grandifolia var. grandifolia

21787 grandifolia var. grandifolia

25768 grandifolia var. grandifolia

26107 grandifolia var. obtusa

QUESTEL, A.

1117 grandifolia var. grandifolia

2929 grandifolia var. grandifolia

RAMSAMMY, J. R.

88 grandifolia var. obtusa

RIDOUTT, C. A.

12975 speciosa

RODRIGUEZ, G.

394 karsteniana

ROIG, J. and EDWARDS*

6429 horrida

ROPER, E. W.

s.n. (GH) horrida

RUSBY, H. H.

149 speciosa

3020 speciosa

RUSBY, H. H. and R. W. SQUIRES

1 10 spectabilis var. spectabilis

SAGOT, P.

873 cruciata

SALVIN, O.

s.n. (GH) decurrens

s.n. (K) decurrens

s.n. (US) decurrens

SARGENT, F. H.

332 horrida

536 horrida

SCAMMAN, E.

5881 choricarpa

6520 /jomrfa

6996 choricarpa

7000 mutica var. mutica

7575 mutica var. mutica

7576 mutica var. mutica

7577 mutica var. mutica

7578 mutica var. mutica

7579 mutica var. mutica

8117 horrida

8143 grandifolia var. grandifolia

SCAMMAN, E. and L. R. HOLDRIDGE

7879 mutica var. contigua

7881 mutica var. mutica

7882 mutica var. mutica

SCHULTES, R. E. and B. P. REKO

678 decurrens

SCHUNKE, C.

52 speciosa

82 speciosa

879 speciosa

A- 143 speciosa

SCHUNKE, J.

4744 speciosa

SEIFRIZ, W.

3 spectabilis var. spectabilis

4 consimilis

13 spectabilis var. spectabilis

277 horrida

94

INDEX TO COLLECTORS' NUMBERS

SEIFRIZ, W. (continued)

572 horrida

SENN, H. A.

230 horrida

387 horrida

SHAFER, J. A.

283 grandifolia var. grandifolia

3270 horrida

4456 horrida

8825 horrida

8950 horrida

SHERRING, R. V.

74 grandifolia var. obtusa

163 grandifolia var. obtusa

164 .grandifolia var. obtusa

174 grandifolia var. obtusa

SIBBER*

(Fl. Martin 375)

grandifolia var. grandifolia

SlNTENIS, P.

417 horrida

1507 horrida

2490B horrida

4168 horrida

6088 horrida

SKUTCH, A. F.

2265 choricarpa

3262 mutica var. mutica

5307 choricarpa

SMITH, A. C.

3584 spectabilis var. spectabilis

3585 spectabilis var. spectabilis

10074 spectabilis var. spectabilis

10516 grandifolia var. grandifolia

SMITH, H. H.

2596 /zomWo

SMITH, H. H. and G. W.

849 grandifolia var. obtusa

854 grandifolia var. obtusa

1715 grandifolia var. obtusa

SMITH, L. B.

2077 uleana var. uleana

SNEIDERN, K. VON

5199 quitensis

A473 quitensis

SODIRO, A.

s.n. (MO) quitensis

s.n. (US) quitensis

SOEJARTO, D. D.

1571 singularis

SOUKUP, J.

1069 speciosa

SPRUCE, R.

3943 horrida

4730A horrida

5364 uleana var. abitaguensis

5365 ewanii

STAHEL and GONGERIJP*

338 spectabilis var. spectabilis

STANDLEY, P. C.

39349 mutica var. mutica

STANDLEY, P. C. and R. TORRES

51250 mutica var. grandis

STANDLEY, P. C. and J. VALERIC

50216 mutica var. mutica

STEHLE, H.

98 grandifolia var. grandifolia

102 grandifolia var. grandifolia

1221 grandifolia var. grandifolia

1774 grandifolia var. grandifolia

STEHLE, H. and M.

3281 grandifolia var. grandifolia

3294 grandifolia var. grandifolia

3422 grandifolia var. grandifolia

4605 grandifolia var. grandifolia

4703 grandifolia var. grandifolia

STEYERMARK, J. A.

61207 spectabilis var. spectabilis

62042 amabilis

89525 spectabilis var. spectabilis

94886 amabilis

94899 consimilis

STEYERMARK, J. A. and G. AGOSTINI

91093 karsteniana

91205 amabilis

STEYERMARK, J. A. and S. NILSSON

218 spectabilis var. spectabilis

STEYERMARK, J. A. and M. RABE

96260 consimilis

STEYERMARK, J. A. et al.

92924 spectabilis var. spectabilis

STOPPERS, A. L.

4203 grandifolia var. grandifolia

4206 grandifolia var. grandifolia

4210 grandifolia var. grandifolia

STORK, H. E.

2580 mutica var. grandis

2582 mutica var. mutica

4756 mutica var. mutica

TAMAYO, F.

154 horrida

2543 horrida

TATE, G. H. M.

424 speciosa

1162 speciosa

TONDUZ, A.

12532 mutica var. mutica

TRIANA, J.

s.n. (1853) quitensis

TRYON, R. M. and A. F.

5260 speciosa

6871 uleana var. uleana

6993 mutica var. mutica

7019 mutica var. mutica

7036 mutica var. grandis

TSCHUDY, R. H. and B. D.

42 horrida

99 karsteniana

132 karsteirana

TURCKHEIM, H. VON

2707 (2717?)

INDEX TO COLLECTORS' NUMBERS

95

TURNER, F. T.

s.n. (US) grandifolia var.

grandifolia

TYSON, E. L. et al.

2452 cocleana

ULE, E.

s.n. (R) uleana var. uleana

UNDERWOOD, L. M.

86 horrida

1548 horrida

2024 horrida

3479 horrida

UNDERWOOD, L. M. and F. S. EARLE

626 horrida

1056 horrida

UNDERWOOD, L. M. and R. F. GRIGGS

71 horrida

269 horrida

273 horrida

VALEUR, E. J.

350 horrida

1018 horrida

VARGAS, C.

11269 horrida

1 1294 speciosa

11714 speciosa

14584 speciosa

14653 speciosa

16073 speciosa

16275 speciosa

16502 speciosa

17738 speciosa

18863 speciosa

VAUTHIER*

586 uleana var. uleana

WACKET, M.

90 uleana var. uleana

303 uleana var. uleana

WATT, D,

110 horrida

WEBSTER, G. L. et al.

9148 grandifolia var. grandifolia

WENT, F. A. F. C.

1018 spectabilis var. spectabilis

WHITE, R. A. and T. W. LUCANSKY

196818 mutica var. mutica

196840 mutica var. chiricana

196852 mutica var. mutica

196854 mutica var. mutica

196857 mutica var. grandis

1968108 mutica var. mutica

1968139 mutica var. mutica

1968168 mutica var. grandis

WIGGINS, I. L.

10930 quitensis

WILBUR, R. L. et al.

7791 grandifolia var. grandifolia

8142 grandifolia var. grandifolia

8185 grandifolia var. grandifolia

10192 mutica var. grandis

10492 mutica var. mutica

10624 mutica var. mutica

10625 mutica var. grandis

11111 cocleana

WILLDENOW, K. L.

20167 grandifolia var. grandifolia

20168 grandifolia var. obtusa

WILLIAMS, LL.

7156 horrida

10758 karsteniana

10786 karsteniana

10985 horrida

WILLIAMS, L. O.

19647 mutica var. grandis

20267 mutica var. mutica

20304 horrida

WILLIAMS, L. O. et al.

28016 mutica var. contigua

28785 choricarpa

29109 mutica var. mutica

WILLIAMS, R. S.

1304 speciosa

WILSON, C. L.

2 1 8 A decurrens

WILSON, K. A. and G. L. WEBSTER

458 horrida

WILSON, P.

290 horrida

WRIGHT, C.

888 horrida

WURDACK, J. J.

1966/zomWc

1991 horrida

2059 nervosa

YUNCKER, T. G.

18061 horrida

18359

INDEX TO NAMES

Recognized species and varieties of Cnemidaria in boldface;

page numbers of descriptions in boldface ; page numbers of illus-

trations in italics.

Actinophlebia, 1, 26, 69

horrida, 26, 43

obtusa, 62

Cnemidaria, 1-3, 6, 26, 27, 80

alatissima, 9, 16, 55, 56

amabilis, 3, 5, 6, 14-16,17, 18, 31,

36,38,42

apiculata, 19, 20, 37, 40

bella, 12, 20, 37, 42, 48, 49

chocoensis, 19, 59, 60

choricarpa, 5, 6, 9, 35, 52, 54, 57,

74,7*

cocleana, 19, 20, 52, 53, 65, 77

conformis, 80

consimilis, 5, 9, 14, 15, 62, 66, 67,

69,76

cruciata, 10, 12, 55, 62, 65, 80

decurrens, 5, 6, 9, 58, 73, 76, 79

dissimilis, 80

ewanii, 5, 11, 15, 70, 76, 78

grandifolia, 14, 16, 26, 36, 37, 42,

43,45,48

var. grandifolia, 16, 37, 43, 46,

79

var. obtusa, 45-47

horrida, 5, 9, 16, 18, 19, 20, 26,

35-37, 42, 50, 61, 79

integrifolia, 80

karsteniana, 5, 18, 75, 76, 77, 79

kohautiana, 46

lindenii, 81

munita, 47

mutica, 5, 12, 18, 20, 32, 3J, 42, 44,

51,58

var. chiricana, 5, 17, 18, 32, 34,

35,58

var. contigua, 15, 33, 34, 37

var. grandis, 33, 35, 36, 58

var. mutica, 5, 9, 15-17, 33, 34,

58,75

nervosa, 5, 18, 20, 54, 65, 76, 77

obtusa, 47

petiolata, 81

quitensis, 9, 12, 15, 20, 54, 54, 58,

71

roraimensis, 71, 72

singularis, 5, 6, 11, 19, 38, 40, 41

speciosa, 1, 3, 5, 6, 11, 15, 26, 51,

59, 68, 71, 76, 81

spectabilis, 5, 9, 21, 50, 52, 61, 65,

66,71,73,80

var. colombiensis, 12, 55, 63, 64

var. spectabilis, 10, 14, 15, 19,

20, 48, 61, 76, 79

tryoniana, 5, 6, 11, 19, 20, 38,

55,40,51

uleana, 6, 15, 18, 19, 38, 50

var. abitaguensis, 38, 50, 51

var. uleana, 17, 50, 51, 78

Cnemidopteris, 26

Cormophyllum horridum, 43

Cyathea, 1,3,5, 14,21,27,80

abitaguensis, 51

ameristoneura, 80

andicola, 54

antillana, 47

apiculata, 42

96

INDEX

97

Cyathea

aristata, 42

bella, 49

chiricana, 36

choricarpa, 57

commutata, 43

conformis, 3, 80

contigua, 37

decurrens, 74

decurrentiloba, 74

dissimilis, 80

elegantissima, 74

ewanii, 70

grandifolia, 46

grandis, 36

guatemalensis, 74

hombersleyii, 81

horrida, 42

imrayana, 46

insignis, 46

integrifolia, 3, 50, 80

karsteniana, 75

kohautiana, 46

leprieurii, 65

liebmanii, 74

lucida, 74

marginalis, 80

maxonii, 51

mexicana, 74

multiflora, 13, 21

munita, 47

mutica, 34

obtusa,.47

var. bullata, 47

var. kohautiana, 47

panamensis, 3, 80

petiolata, 81

pittieri, 34

quitensis, 54

roraimensis, 71

rudis, 36

speciosa, 3, 21, 50, 68, 69, 81

spectabilis

var. longipinna, 62

var. trinitensis, 62

subarachnoidea, 35

subarborescens, 51

subglabra, 35

subincisa, 68

wilsonii, 81

woronovii, 3, 81

Hemistegia, 1, 26, 69

ameristoneura, 77

decurrens, 73

elegantissima, 73

grandifolia, 46

horrida, 43

insignis, 46

kohautiana, 26, 46

lucida, 73

marginalis, 81

mexicana, 73

monilifera, 80

munita, 47

obtusa, 47

repanda, 43

speciosa, 68

spectabilis, 62

willdenowii, 46

Hemitelia, 1,26, 27

abitaguensis, 51

acuminata, 43

amabilis, 31

ameristoneura, 77

apiculata, 40, 42

arachnoidea, 34, 35

bella, 48

bullata, 47

chiricana, 35

chiriquana, 35

choricarpa, 57

commutata, 43

conformis, 80

contigua, 37

cruciata, 65

decurrens, 73, 75

dissimilis, 80

grandifolia, 45, 46

grandis, 36

guatemalensis, 73

hombersleyii, 81

hookeri, 43

hookeriana, 43

horrida, 43

var. heterosora, 34

var. imrayana, 43, 46

imrayana 46, 47

insignis, 46

98

INDEX

Hemistegia

integrifolia, 80

karsteniana, 75, 76

klotzschiana, 62, 76

kohautiana, 45, 46

leprieurii, 65

lindenii, 81

lucida, 73

marginalis, 80

maxonii, 51

mexicana, 74, 75

monilifera, 46, 80

munita, 47

mutica, 34

nervosa, 77

obtusa, 45, 47, 48, 62

var. bullata, 47

var. kohautiana, 46

petiolata, 81

pittieri,34,35,58

quitensis, 54, 55

roraimensis, 71

rudis, 36

serrata, 46

speciosa.21,69,70,81

spectabilis, 61

var. longipinna, 62

var. trinitensis, 62

subglabra, 34

subincisa, 68

uleana, 51

venosa, 81

wilsonii, 81

woronovii, 81

Microstegnus, 1, 26

grandifolius, 26, 46

Polypodium horridum, 42

Trichipteris, 3, 27

williamsii, 3

Publication 1194

UNIVERSITY OF ILLINOIS-URBANA